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Os03g0227700

2014-05-27T13:32:41Z

LQ: /* Expression */

Please input one-sentence summary here.
The gene OsDWARF4 was found in young elongating tissues.
==Annotated Information==

Brassinosteroids influence both plant height and leaf erectness in rice<ref name="ref1" />. Arabidopsis thaliana BRASSINOSTEROID INSENSITIVE1 (bri1) mutants, which are deficient in the brassinosteroid receptor, have a severe dwarf phenotype<ref name="ref2" />. The rice mutants d61-1 and d61-2, which are weak mutant alleles of OsBRI1, a homolog of A. thaliana BRI1, have both a semi-dwarf phenotype and more erect leaves<ref name="ref3" />. In rice, an erect leaf phenotype has also been associated with loss-of-function, brassinosteroid-deficient mutants such as brassinosteroid-deficient dwarf1 (brd1), which is defective in OsDWARF (the homolog of tomato CYP85A1/DWARF<ref name="ref4" />), and ebisu dwarf (d2), which is defective in CYP90D2/D2 (ref.<ref name="ref5" />). However, the grain yield of d61, brd1 and d2 mutants is decreased because of morphological alterations in their reproductive development.

===Function===

New cultivars with very erect leaves, which increase light capture for photosynthesis and nitrogen storage for grain filling, may have increased grain yields.The erect leaf phenotype of a rice brassinosteroid-deficient mutant, osdwarf4-1, is associated with enhanced grain yields under conditions of dense planting, even without extra fertilizer. Molecular and biochemical studies reveal that two different cytochrome P450s, CYP90B2/OsDWARF4 and CYP724B1/D11, function redundantly in C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis. Therefore, despite the central role of brassinosteroids in plant growth and development, mutation of OsDWARF4 alone causes only limited defects in brassinosteroid biosynthesis and plant morphology<ref name="ref6" />.

===Expression===
The affected gene, OsDWARF4, was identified by characterizing the site of insertion of the Tos17 retrotransposon.

----Molecular characterization of OsDWARF4 and OsDWARF4L1/D11

OsDWARF4 is a homolog of A. thaliana DWARF4, which encodes a cytochrome P450, CYP90B1 (Fig. 1a). CYP90B1/DWARF4 catalyzes C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis<ref name="ref7" />. Only one DWARF4 homolog was identified in the rice genome. OsDWARF4 is located on the short arm of chromosome 3 (35 cM). Its predicted open reading frame consists of eight exons, and encodes a protein of 502 amino acids (Fig. 1b,c). OsDWARF4 is most closely related to DWARF4 (66.3% amino-acid identity; Fig. 1a). The structure of OsDWARF4 is similar to that of DWARF4 throughout its length: five domains found in cytochrome P450s—proline rich, A (also referred to as dioxygen binding), B (steroid binding), C and heme binding—are highly conserved (Fig. 1c). In wild-type rice, OsDWARF4 transcripts are most abundant in leaf blades and roots, but are also found in all other plant parts tested (Fig. 1d). Transcript levels were decreased by exogenous brassinolide treatment, and increased in brassinosteroid-insensitive d61-3 and brassinosteroid-deficient brd1-1 (Fig. 1e). These results suggest that OsDWARF4 expression is feedback regulated, as is that of A. thaliana brassinosteroid-biosynthesis genes<ref name="ref8" />.
OsDWARF4L1 was identified as an OsDWARF4-like gene by in silico screening of rice DNA databases. OsDWARF4L1 is located on the long arm of chromosome 4 (70.9 cM). A recent study demonstrated that a rice semi-dwarf and short-grain mutant, d11, has a loss-of-function mutation in the OsDWARF4L1 gene<ref name="ref9" />. OsDWARF4L1/D11 shows the highest homology with A. thaliana CYP724A1, and is closely related to rice OsDWARF4 and A. thaliana DWARF4 (47.4%, 37.7% and 41.7% amino-acid identities, respectively; Fig. 1a). In wild-type rice, OsDWARF4L1/D11 transcripts were most abundant in vegetative shoot apices, leaf sheaths and roots, and found at more moderate levels in other organs (Fig. 1d). They were decreased by exogenous brassinolide treatment, and increased in d61-3 and brd1-1 (Fig. 1e). These results indicate that OsDWARF4L1/D11 expression is regulated by a feedback mechanism, as is that of OsDWARF4.
[[File:Figure_1_-_Molecular_characterization_of_OsDWARF4_and_OsDWARF4L1-D11..jpg|right|thumb|500px|'''Figure 1.''' ''Molecular characterization of OsDWARF4 and OsDWARF4L1/D11 (a) Phylogenetic relationships among brassinosteroid-biosynthetic cytochrome P450s of rice, A. thaliana and tomato. Bar, 0.1 amino-acid substitutions per site. (b) Genomic organization of the OsDWARF4 gene. Boxes represent exons, and lines separating boxes represent introns. The arrowhead indicates the site of Tos17 insertion in osdwarf4-1. (c) Relative positions of the major domains in OsDWARF4. All of the major domains found in the cytochrome P450 superfamily are conserved in OsDWARF4. The arrowhead indicates the site of Tos17 insertion in osdwarf4-1. (d) Expression of OsDWARF4 and OsDWARF4L1/D11 in various organs of wild-type rice. The gene encoding Histone H3 was used as a control. (e) Feedback regulation of OsDWARF4 and OsDWARF4L1/D11 in brassinolide-treated wild-type plants (left) and brassinosteroid-related mutants (right). Values under each panel indicate relative amounts of each transcript. (f) Function of recombinant CYP90B2/OsDWARF4 and CYP724B1/D11 proteins. Campesterol (CR) was incubated with each recombinant protein and analyzed by GC-MS. Inset shows that the mass spectrum of the trimethylsilyl ester of each metabolite coincided with that of the trimethylsilyl ether of synthesized (22S)-22-hydroxycampesterol (22-OHCR).]]
----Phenotypes of rice mutants defective in brassinosteroid biosynthesis

In osdwarf4-1, Tos17 insertion into exon 6 causes a premature termination in domain C (arrowhead in Fig. 1b,c). Because a premature stop codon before the heme-binding domain caused loss-of-function of A. thaliana DWARF4 (ref.<ref name="ref7" />), we consider osdwarf4-1 to be a null allele. Unexpectedly, osdwarf4-1 plants show only weak morphological phenotypes, such as a slightly dwarfed stature and more erect leaves (Fig. 2a,b). Both of these phenotypes were complemented by the introduction of an entire OsDWARF4 gene (data not shown). These results seem inconsistent with the fact that rice d61-3 and brd1-1, which are loss-of-function mutants of OsBRI1 and OsDWARF respectively, display severe dwarfing and have malformed leaves (for example, see brd1-1 in Fig. 2a), and suggest functional redundancy in brassinosteroid biosynthesis in rice.
Because null alleles of d11, such as d11-4, also show a weak phenotype<ref name="ref9" />(Fig. 2a),OsDWARF4L1/D11 and OsDWARF4 may function redundantly. To confirm this possibility, researcher produced double mutants by crossing homozygous osdwarf4-1 and d11-4 plants. Among the F2 population, 34 of 576 plants showed severe dwarfing and malformed leaves with tortuous leaf blades, indistinguishable from the brassinosteroid-deficient brd1-1 phenotype (Fig. 2a,c,d). This ratio fits the theoretical 9:3:3:1 (wild-type:osdwarf4-1:d11-4:osdwarf4-1/d11-4) segregation predicted by mendelian inheritance. PCR and sequence analyses confirmed that these plants were osdwarf4-1/d11-4 double mutants (data not shown).
Endogenous levels of brassinosteroid intermediates in brd1 clearly indicate that OsDWARF catalyzes C-6 oxidation<ref name="ref4" />. However, quantification in d2 could not determine the step(s) catalyzed by CYP90D2/D2, because CYP90D2/D2 functions redundantly with another gene, CYP90D3, and loss-of-function of CYP90D2/D2 have caused only limited defects in brassinosteroid biosynthesis<ref name="ref5" />. Similarly, endogenous levels of brassinosteroid intermediates were obviously changed in the osdwarf4-1/d11-4 double mutant relative to osdwarf4-1 or d11 single mutants. Although brassinolide was not detected in either the wild-type or osdwarf4-1/d11-4, another bioactive brassinosteroid, castasterone (CS), was detected in the wild-type but not in osdwarf4-1/d11-4, confirming that osdwarf4-1/d11-4 is brassinosteroid-deficient (Fig. 2e). Levels of the other five intermediates downstream of C-22 hydroxylation—6-deoxocathasterone (6-DeoxoCT), 6-deoxoteasterone (6-DeoxoTE), 3-dehydro-6-deoxoteasterone (6-Deoxo3DT), 6-deoxotyphasterol (6-DeoxoTY) and 6-deoxocastasterone (6-DeoxoCS)—were also greatly reduced in osdwarf4-1/d11-4 (Fig. 2e). This indicates that both proteins are involved in C-22 hydroxylation. It is unclear why the upstream compound, campestanol, did not accumulate in osdwarf4-1/d11-4. Because A. thaliana CYP90B1/DWARF4 also catalyzes C-22 hydroxylation of other upstream compounds such as campesterol and (24R)-ergost-4-en-3-one<ref name="ref10" />, it is possible that campestanol is not the major substrate for CYP90B2/OsDWARF4 and CYP724B1/D11 in rice.
[[File:Initpintu_副本.jpg|right|thumb|500px|'''Figure 2.''' ''Phenotypes of rice mutants defective in brassinosteroid biosynthesis.(a) Comparison of gross morphology between the wild-type, single mutants (osdwarf4-1, d11-4, brd1-1), and a double mutant (osdwarf4-1/d11-4). Bar, 30 cm. (b) Erect leaf phenotype of osdwarf4-1. The degree of bending between leaf blade and sheath of osdwarf4-1 (right) is less than that of the wild-type (left). lb, leaf blade; ls, leaf sheath. Bar, 3 cm. (c,d) Close-up views of the osdwarf4-1/d11-4 double mutant (c) and brd1-1 (d). Bars, 5 cm. (e) Endogenous contents of brassinosteroid intermediates in the wild-type and osdwarf4-1/d11-4 double mutant. CN, campestanol; 6-DeoxoCT, 6-deoxocathasterone; 6-DeoxoTE, 6-deoxoteasterone; 6-Deoxo3DT, 3-dehydro-6-deoxoteasterone; 6-DeoxoTY, 6-deoxotyphasterol; 6-DeoxoCS, 6-deoxocastasterone; CS, castasterone; BL, brassinolide.]]
----Characterization of a field-grown osdwarf4-1 mutant

Because the harvest index (grain/(grain plus straw)) of rice is about 0.5, further increases in yield potential will have to involve increases in crop biomass driven by more net photosynthesis<ref name="ref11" />. In rice, the contribution of lower leaves to photosynthesis is significant, even though their photosynthetic capacity is considerably less than that of upper leaves<ref name="ref12" />. Erect leaves allow greater penetration of light to lower leaves, thereby optimizing canopy photosynthesis<ref name="ref13" />. Therefore, the most likely explanation for the increase in above-ground biomass in osdwarf4-1 plots under dense planting conditions (more than 1.2 times that in wild-type under conventional planting condition) is that the shade of the upper leaves is minimized, and the lower leaves receive more light to drive higher rates of photosynthesis (Fig. 3 and Supplementary Table 2 online). No difference was observed in the heading date (flowering time) and the grain-filling period (time to maturity) between wild-type and osdwarf4-1 plants (data not shown). In the wild-type, the above-ground biomass was increased in response to higher fertilizer application under conventional planting (22.2 plants m-2), but was significantly decreased under dense planting with doubled fertilizer application owing to reduced photosynthesis caused by lodging (Fig. 3a). In contrast, the above-ground biomass of osdwarf4-1 was significantly (P < 0.01) increased by dense planting at each fertilizer rate. Consequently, the above-ground biomass in osdwarf4-1 plots under dense planting with standard and a 1.5-fold increase in fertilizer application was approx1.34 and 1.52 times that in wild-type plots under conventional cultivation conditions. The increased biomass depended mainly on accelerated formation of tillers, which have the ability to generate a panicle (panicle number, Fig. 3b). Both total and fertile grain numbers tended to increase in densely planted osdwarf4-1 plots (Fig. 3c), resulting in significantly increased grain yield even when the fertilizer application was not increased (for example, the osdwarf4-1 yield was 6.19 t ha-1 under dense planting with standard fertilizer application whereas the wild-type yield was 4.69 t ha-1 under conventional conditions; Fig. 3d). However, the increase in grain yield was not as much as that in above-ground biomass, because the harvest index of osdwarf4-1 (0.41) was slightly lower than that of the wild-type (0.44). A similar field test gave comparable results.
[[File:搜狗截图14年05月24日2050_11_副本.png|left|thumb|500px|'''Figure 3.''' ''Characterization of a field-grown osdwarf4-1 mutant.(a) Comparison of biomass production between the wild-type and osdwarf4-1. White and black bars indicate above-ground and panicle dry weight, respectively. (b) Comparison of panicle number (white bar) and panicle length (black bar). (c) Comparison of total (white bar) and fertile (black bar) grain number. (d) Comparison of estimated grain yields between wild-type (white bar) and osdwarf4-1 (black bar). W, wild-type, M, osdwarf4-1. Density indicates conventional planting (22.2 plants m-2) or dense planting (44.4 plants m-2). Fertilizer indicates the level of nitrogen fertilizer. Conventional condition is times 1 (6 g m-2), and two increased conditions are times 1.5 and times 2 (9 and 12 g m-2 respectively). Lowercase letters indicate significant differences at the level of P < 0.01 within a parameter (Tukey's Honest Significant Difference test).]]

----Overexpression of OsDWARF4

Overexpression of DWARF in transgenic tomato increases plant height<ref name="ref14" />, and overexpression of DWARF4 in transgenic A. thaliana and tobacco increases vegetative growth and seed yield<ref name="ref15" />. In rice, however, the role of brassinosteroid in regulating leaf angle has been known for over 40 years<ref name="ref16" />,<ref name="ref17" />. Whereas brassinosteroid deficiency increased the erectness of leaves, the opposite phenotype (increased leaf angle) was observed in transgenic rice overexpressing OsDWARF4 (27.6° versus 10.0° in the wild-type, P < 0.01), and was inherited in their progeny (data not shown). Because inclined leaves are unfavorable for photosynthesis when plants are grown in a group, we have to carefully evaluate whether the overexpression of a brassinosteroid-biosynthesis gene in transgenic rice increases grain yield as is the case in transgenic plants overexpressing DWARF4.

===Conclusion===

More erect leaves improve not only light distribution but also the size and use of leaf nitrogen reservoirs for grain growth<ref name="ref13" />. The erect leaf phenotype caused by brassinosteroid deficiency improved biomass production under field conditions, resulting in increased grain yield without the negative environmental effects associated with fertilizers. Recently identified rice homologs of A. thaliana BAS1 that encodes the brassinosteroid catabolic enzyme, CYP734A1 (ref.<ref name="ref18" />). Overexpression of these rice homologs in transgenic rice successfully induced the brassinosteroid-deficient phenotypes (unpublished results). Although ideally, expression of these genes should be restricted to particular tissues and organs such as leaf laminae, it should be possible to generate erect leaf varieties by the introduction of a single transgene. In conclusion, engineering more erect leaves modulating brassinosteroid metabolism can be combined with dense planting to improve rice grain yields.

==Labs working on this gene==
Please input related labs here.

==References==
<references>
<ref name="ref1">Sakamoto, T. & Matsuoka, M. Generating high-yielding varieties by genetic manipulation of plant architecture. Curr. Opin. Biotech. 15, 144–147 (2004).</ref>
<ref name="ref2">Li, J. & Chory, J. A putative leucine-rich repeat receptor kinase involved in brassinosteroid signal transduction. Cell 90, 929–938 (1997).</ref>
<ref name="ref3">Yamamuro, C. et al. Loss of function of a rice brassinosteroid insensitive1 homolog prevents internode elongation and bending of the lamina joint. Plant Cell 12, 1591–1606 (2000).</ref>

<ref name="ref4">Hong, Z. et al. Loss-of-function of a rice brassinosteroid biosynthetic enzyme, C-6 oxidase, prevents the organized arrangement and polar elongation of cells in the leaves and stem. Plant J. 32, 495–508 (2002).</ref>
<ref name="ref5">Hong, Z. et al. A rice brassinosteroid-deficient mutant, ebisu dwarf (d2), is caused by a loss of function of a new member of cytochrome P450. Plant Cell 15, 2900–2910 (2003).</ref>
<ref name="ref6">Sakamoto T;Morinaka Y;Ohnishi T;Sunohara H;Fujioka S;Ueguchi Tanaka M;Mizutani M;Sakata K;Takatsuto S;Yoshida S;Tanaka H;Kitano H;Matsuoka M Erect leaves caused by brassinosteroid deficiency increase biomass production and grain yield in rice Nature Biotechnology, 2006, 24(1): 105-109.</ref>
<ref name="ref7">Choe, S. et al. The DWF4 gene of Arabidopsis encodes a cytochrome P450 that mediates multiple 22alpha-hydroxylation steps in brassinosteroid biosynthesis. Plant Cell 10, 231–243 (1998).</ref>
<ref name="ref8">Bancos, S. et al. Regulation of transcript levels of the Arabidopsis cytochrome P450 genes involved in brassinosteroid biosynthesis. Plant Physiol. 130, 504–513 (2002).</ref>
<ref name="ref9">Tanabe, S. et al. A novel cytochrome P450 is implicated in brassinosteroid biosynthesis via the characterization of a rice dwarf mutant, dwarf11, with reduced seed length. Plant Cell 17, 776–790 (2005).</ref>
<ref name="ref10">Fujioka, S., Takatsuto, S. & Yoshida, S. An early C-22 oxidation branch in brassinosteroid biosynthetic pathway. Plant Physiol. 130, 930–939 (2002).</ref>
<ref name="ref11">Mann, C.C. Crop scientists seek a new revolution. Science 283, 310–314 (1999).</ref>
<ref name="ref12">Horton, P. Prospects for crop improvement through the genetic manipulation of photosynthesis: morphological and biochemical aspects of light capture. J. Exp. Bot. 51, 475–485 (2000).</ref>
<ref name="ref13">Sinclair, T.R. & Sheehy, J.E. Erect leaves and photosynthesis in rice. Science 283, 1455 (1999).</ref>
<ref name="ref14">Bishop, G.J. et al. The tomato DWARF enzyme catalyses C-6 oxidation in brassinosteroid biosynthesis. Proc. Natl. Acad. Sci. USA 96, 1761–1766 (1999).</ref>
<ref name="ref15">Choe, S. et al. Overexpression of DWARF4 in the brassinosteroid biosynthetic pathway results in increased vegetative growth and seed yield in Arabidopsis. Plant J. 26, 573–582 (2001).</ref>
<ref name="ref16">Maeda, E. Rate of lamina inclination in excised rice leaves. Physiol. Plant. 18, 813–827 (1965).</ref>
<ref name="ref17">Wada, K. et al. Brassinolide and homobrassinolide promotion of lamina inclination of rice seedlings. Plant Cell Physiol. 22, 323–325 (1981).</ref>
<ref name="ref18">Neff, M.M. et al. BAS1: A gene regulating brassinosteroid levels and light responsiveness in Arabidopsis. Proc. Natl. Acad. Sci. USA 96, 15316–15323 (1999).</ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os03g0227700|
Description = Cytochrome P450 family protein|
Version = NM_001055982.2 GI:297600576 GeneID:4332134|
Length = 6529 bp|
Definition = Oryza sativa Japonica Group Os03g0227700, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 3|Chromosome 3]]|
AP = Chromosome 3:6765109..6771637|
CDS = 6765109..6765374,6765543..6765867,6765971..6766123,6768937..6769188,6769337..6769429 ,6770222..6770300,6770398..6770504,6771392..6771637|
GCID = <gbrowseImage1>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</cdnaseq>|
AA = <aaseq>MAAMMASITSELLFFLPFILLALLTFYTTTVAKCHGGHWWRGGT TPAKRKRMNLPPGAAGWPLVGETFGYLRAHPATSVGRFMEQHIARYGKIYRSSLFGER TVVSADAGLNRYILQNEGRLFECSYPRSIGGILGKWSMLVLVGDPHREMRAISLNFLS SVRLRAVLLPEVERHTLLVLRAWPPSSTFSAQHQAKKFTFNLMAKNIMSMDPGEEETE RLRREYITFMKGVVSAPLNLPGTPYWKALKSRAAILGVIERKMEERVEKLSKEDASVE QDDLLGWALKQSNLSKEQILDLLLSLLFAGHETSSMALALAIFFLEGCPKAVQELREE HLGIARRQRLRGECKLSWEDYKEMVFTQCVINETLRLGNVVRFLHRKVIKDVHYKGYD IPSGWKILPVLAAVHLDSSLYEDPQRFNPWRWKSSGSSGGLAQSSSFMPYGGGTRLCA GSELAKLEMAVFLHHLVLNFRWELAEPDQAFVFPFVDFPKGLPIRVHRIAQDDEQE</aaseq>|
DNA = <dnaseqindica>1..266#435..759#863..1015#3829..4080#4229..4321#5114..5192#5290..5396#6284..6529#atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtcggtggtttcattactcggtacacgcacacagctatataagcacgcgcactgctcgcctcgccgtgaagcaagcttagctagtagctaggatatcgatcgatcggtaataataggctgacggcgagctggggggtggcgtgcggttgcgtgcgtgcgtgcatgcaggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaaggtacaacacacatcgtcctccattaagttgagcatgcatatatcaatcagctcgctctagcgatcaatatcaatgaaacgcgcgtttgcgtgcgtgcgtgcagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaaggtacgtactcgtagctactagctactccctccgtttcgaaatgtttgacgccgttgactttttatcacatatttgaccgttcgttttatttaaaaaaattaagtaattattaattattttcctatcatttgattcattgttaaatatatttttatgtaggcatataattttacatatctcacaaaagtttttaaataagacgaacggtcaaacatttgctaaaaagtcaacggtgtcaaatatttcgaaacggaggaagtagctagctaccaccccacccgtgcggattgatcgatcgatcgatgtccctctctgcctacacacaccctgcgccccggccgtacgcgcgcaaagtcgtgcgcgcatgagttttactccttccgtttcaaaaaaaaaacgacctagacaagtaatatgatgtaacctagaacaacgaatctatacaggagtatatccaaattcattgtcctagattacatcatatctttgtctatgttgccttttttaggaggagagagtactttacatatgagttatgaccggccggccaggttcctttcttcccctattccgtatgcatactataagcagctgcatataggcatatactactatacccagcttagctaaagcaaacaacttgcaacgacgatcatgtgcaggtccctgttgacagaggttagagatgtaaaccatttttgttgttttcattatctaaagagaaaacaagttactacgacgatcgagatgtacgcaaggggatcatatttatattggagaggagagagagatgatttgcgacttgtgagtgataagtgaatatatatactatatatttgtgtaggagtgacgatgacttccctgtcaatgttttgtttggtatgcacacggggggaaaagagtttagagaaaggaattggctagagagggccaggtttagtttctttaatttttgtttggtggttttgtgtgtagctagtccgcatgcatgatatatcatatatatgcatggcccatcattagatatatatacatgacggcagtgcacttattatatatatgtaggggcatatacaaggtgtatgtgtttagaattgtggatggtccaggaattgttgcagctagttctatactagcatagtgaattagtgatccatatcgagatcaggtccaatcggtcagctcgcacagcgggtcttttagtacgatttgttcggacaaattatcttgcctttgagataatcaacataggattgaaactgcatactagcaatagaccatagctatttggcaatatagaaagctaaaaaatgttgattgaaaggtgcgtgcatgcatgctagcagtatgtcttttaggccaggtccagtttttttttttttctcaaaggcagttgcttaaggtgaaattctagcagaggtcatatgaatatatttattatcttccaaacatgagccaaaacctatatatagttaactcaaaaaaccatgcattggtacttcttctgagtatgcatcctggggactgatcgatcatgcagattcttttctttctttttccctggggacgtactgatcatcactcatgtgagcataaaatgttgcctactttttactgcatgtgaggtcacactcaaatgatctttattgattgcctcagcatgcatgctgctagctttcacgatctctctccgatacacacaccctgtgctgtgctcaacagcatgcaacctggcaggtggaatttttaatctgttggaaagtaaatgaccaaccttcataatttggcatgattatatgctggggaatctatcatttatcatgattcaaccgctgaccaaactctttgccacacacccgaaccacattaatttcgttaagcggatgattagccagcaaccttcacagtttggttttaattagttaatgaaagggtgcttgtagttttattagctgaagagacactgcagtactcctactgagcgttctccttaattgggaaatagatctctgcagtactatcctttaactttaagtgaataacatgtagctaggtctcacacttatcaggcccacacatcagtgatccaagtcaaatggcaatacaccataggagattcttccactaattggctaacaaaaataaactgtgatcactgatcacagctgctaacactataacacattaatttgcatgtctcatgaaatgccagatgagttggattaattatctatcttatgttcagtgccgaggaatatacaaggaaacatcattggtatagttgtggagttctgagtgcattatctcacccagtacggtcaaattaatcctagtctccgtaaatatggttgtgtgcactcttagttaattagtgtagacccggaaattatactttgtcgagaatgttgttatccgtgtgttaagtttcaagtaccctgcaggcctataactaaacactgccaacatggatgtttaagaatgaattatacatacagcgcacttggaaaagaaaaggggtagctacccagctgaagaacacctctctctcatttgaaggtcgacacaatatgtgtcttttctatatcgaaaggaaacgacattttttctcttcaaaacagaaagctatgacaagagacataatcctagactagtaggcggttaggcaacaccatatctgaatttgataggtgtcccacatattttttttcttttcctgataagaaggaacttttcaccgtgagcttatttttttctgttctatgtcgacattagttccatgatgtttactgtgtatttttactgttcccctgcagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggtacagagaattacagattcgaaattaacatctactgtcactagctcagtgctacgtttaactgtagagcaacaattgcagaaagcgaaggcatcagaaatgacatgggaatcgattagttacactcacactgacgcgtgacatgcaggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtaaggccccccctactttgttccttccgatccctcctccctaaacgctactaatataaatcggtgcaaaggctttgtgctagcgtagctcttgctgttttgtttggtcacttgcgtagagagatagcaacttcccgaaatgtcatgtacgaatggagactgaaagctctgcgaaataattgccgggccacgtactacagtacttacgctccctctcttcagtttggatcgatcgagctaagaaggcgcacgcatgtgtgcattgatcgatagagtcgatctgttctgccttgtaagctagccagctgaggttctttcgccttggaagtttctacagggtgaggcccatcgtcttgcaacgatccaaatgccatgacaagtacctgttgttgtaggcttccagtagcacggcctgcccccaagaaggtatctgaatcttctgcattgctgttcagacggcagagcagaacatgccgaaataccattccagtcagttgcagtatatgtgtgtggtggtatcagataaaggagcagagtaatttgaggccttagaaattcagatgagttgctcgacaattgtcgcc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Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001055982.2 RefSeq:Os03g0227700]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 3]]
[[Category:Chromosome 3]]

File:搜狗截图14年05月24日2050 11 副本.png

2014-05-27T13:30:26Z

LQ:

Os03g0227700

2014-05-27T13:29:16Z

LQ: /* Expression */

Please input one-sentence summary here.
The gene OsDWARF4 was found in young elongating tissues.
==Annotated Information==

Brassinosteroids influence both plant height and leaf erectness in rice<ref name="ref1" />. Arabidopsis thaliana BRASSINOSTEROID INSENSITIVE1 (bri1) mutants, which are deficient in the brassinosteroid receptor, have a severe dwarf phenotype<ref name="ref2" />. The rice mutants d61-1 and d61-2, which are weak mutant alleles of OsBRI1, a homolog of A. thaliana BRI1, have both a semi-dwarf phenotype and more erect leaves<ref name="ref3" />. In rice, an erect leaf phenotype has also been associated with loss-of-function, brassinosteroid-deficient mutants such as brassinosteroid-deficient dwarf1 (brd1), which is defective in OsDWARF (the homolog of tomato CYP85A1/DWARF<ref name="ref4" />), and ebisu dwarf (d2), which is defective in CYP90D2/D2 (ref.<ref name="ref5" />). However, the grain yield of d61, brd1 and d2 mutants is decreased because of morphological alterations in their reproductive development.

===Function===

New cultivars with very erect leaves, which increase light capture for photosynthesis and nitrogen storage for grain filling, may have increased grain yields.The erect leaf phenotype of a rice brassinosteroid-deficient mutant, osdwarf4-1, is associated with enhanced grain yields under conditions of dense planting, even without extra fertilizer. Molecular and biochemical studies reveal that two different cytochrome P450s, CYP90B2/OsDWARF4 and CYP724B1/D11, function redundantly in C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis. Therefore, despite the central role of brassinosteroids in plant growth and development, mutation of OsDWARF4 alone causes only limited defects in brassinosteroid biosynthesis and plant morphology<ref name="ref6" />.

===Expression===
The affected gene, OsDWARF4, was identified by characterizing the site of insertion of the Tos17 retrotransposon.

----Molecular characterization of OsDWARF4 and OsDWARF4L1/D11

OsDWARF4 is a homolog of A. thaliana DWARF4, which encodes a cytochrome P450, CYP90B1 (Fig. 1a). CYP90B1/DWARF4 catalyzes C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis<ref name="ref7" />. Only one DWARF4 homolog was identified in the rice genome. OsDWARF4 is located on the short arm of chromosome 3 (35 cM). Its predicted open reading frame consists of eight exons, and encodes a protein of 502 amino acids (Fig. 1b,c). OsDWARF4 is most closely related to DWARF4 (66.3% amino-acid identity; Fig. 1a). The structure of OsDWARF4 is similar to that of DWARF4 throughout its length: five domains found in cytochrome P450s—proline rich, A (also referred to as dioxygen binding), B (steroid binding), C and heme binding—are highly conserved (Fig. 1c). In wild-type rice, OsDWARF4 transcripts are most abundant in leaf blades and roots, but are also found in all other plant parts tested (Fig. 1d). Transcript levels were decreased by exogenous brassinolide treatment, and increased in brassinosteroid-insensitive d61-3 and brassinosteroid-deficient brd1-1 (Fig. 1e). These results suggest that OsDWARF4 expression is feedback regulated, as is that of A. thaliana brassinosteroid-biosynthesis genes<ref name="ref8" />.
OsDWARF4L1 was identified as an OsDWARF4-like gene by in silico screening of rice DNA databases. OsDWARF4L1 is located on the long arm of chromosome 4 (70.9 cM). A recent study demonstrated that a rice semi-dwarf and short-grain mutant, d11, has a loss-of-function mutation in the OsDWARF4L1 gene<ref name="ref9" />. OsDWARF4L1/D11 shows the highest homology with A. thaliana CYP724A1, and is closely related to rice OsDWARF4 and A. thaliana DWARF4 (47.4%, 37.7% and 41.7% amino-acid identities, respectively; Fig. 1a). In wild-type rice, OsDWARF4L1/D11 transcripts were most abundant in vegetative shoot apices, leaf sheaths and roots, and found at more moderate levels in other organs (Fig. 1d). They were decreased by exogenous brassinolide treatment, and increased in d61-3 and brd1-1 (Fig. 1e). These results indicate that OsDWARF4L1/D11 expression is regulated by a feedback mechanism, as is that of OsDWARF4.
[[File:Figure_1_-_Molecular_characterization_of_OsDWARF4_and_OsDWARF4L1-D11..jpg|right|thumb|500px|'''Figure 1.''' ''Molecular characterization of OsDWARF4 and OsDWARF4L1/D11 (a) Phylogenetic relationships among brassinosteroid-biosynthetic cytochrome P450s of rice, A. thaliana and tomato. Bar, 0.1 amino-acid substitutions per site. (b) Genomic organization of the OsDWARF4 gene. Boxes represent exons, and lines separating boxes represent introns. The arrowhead indicates the site of Tos17 insertion in osdwarf4-1. (c) Relative positions of the major domains in OsDWARF4. All of the major domains found in the cytochrome P450 superfamily are conserved in OsDWARF4. The arrowhead indicates the site of Tos17 insertion in osdwarf4-1. (d) Expression of OsDWARF4 and OsDWARF4L1/D11 in various organs of wild-type rice. The gene encoding Histone H3 was used as a control. (e) Feedback regulation of OsDWARF4 and OsDWARF4L1/D11 in brassinolide-treated wild-type plants (left) and brassinosteroid-related mutants (right). Values under each panel indicate relative amounts of each transcript. (f) Function of recombinant CYP90B2/OsDWARF4 and CYP724B1/D11 proteins. Campesterol (CR) was incubated with each recombinant protein and analyzed by GC-MS. Inset shows that the mass spectrum of the trimethylsilyl ester of each metabolite coincided with that of the trimethylsilyl ether of synthesized (22S)-22-hydroxycampesterol (22-OHCR).]]
----Phenotypes of rice mutants defective in brassinosteroid biosynthesis

In osdwarf4-1, Tos17 insertion into exon 6 causes a premature termination in domain C (arrowhead in Fig. 1b,c). Because a premature stop codon before the heme-binding domain caused loss-of-function of A. thaliana DWARF4 (ref.<ref name="ref7" />), we consider osdwarf4-1 to be a null allele. Unexpectedly, osdwarf4-1 plants show only weak morphological phenotypes, such as a slightly dwarfed stature and more erect leaves (Fig. 2a,b). Both of these phenotypes were complemented by the introduction of an entire OsDWARF4 gene (data not shown). These results seem inconsistent with the fact that rice d61-3 and brd1-1, which are loss-of-function mutants of OsBRI1 and OsDWARF respectively, display severe dwarfing and have malformed leaves (for example, see brd1-1 in Fig. 2a), and suggest functional redundancy in brassinosteroid biosynthesis in rice.
Because null alleles of d11, such as d11-4, also show a weak phenotype<ref name="ref9" />(Fig. 2a),OsDWARF4L1/D11 and OsDWARF4 may function redundantly. To confirm this possibility, researcher produced double mutants by crossing homozygous osdwarf4-1 and d11-4 plants. Among the F2 population, 34 of 576 plants showed severe dwarfing and malformed leaves with tortuous leaf blades, indistinguishable from the brassinosteroid-deficient brd1-1 phenotype (Fig. 2a,c,d). This ratio fits the theoretical 9:3:3:1 (wild-type:osdwarf4-1:d11-4:osdwarf4-1/d11-4) segregation predicted by mendelian inheritance. PCR and sequence analyses confirmed that these plants were osdwarf4-1/d11-4 double mutants (data not shown).
Endogenous levels of brassinosteroid intermediates in brd1 clearly indicate that OsDWARF catalyzes C-6 oxidation<ref name="ref4" />. However, quantification in d2 could not determine the step(s) catalyzed by CYP90D2/D2, because CYP90D2/D2 functions redundantly with another gene, CYP90D3, and loss-of-function of CYP90D2/D2 have caused only limited defects in brassinosteroid biosynthesis<ref name="ref5" />. Similarly, endogenous levels of brassinosteroid intermediates were obviously changed in the osdwarf4-1/d11-4 double mutant relative to osdwarf4-1 or d11 single mutants. Although brassinolide was not detected in either the wild-type or osdwarf4-1/d11-4, another bioactive brassinosteroid, castasterone (CS), was detected in the wild-type but not in osdwarf4-1/d11-4, confirming that osdwarf4-1/d11-4 is brassinosteroid-deficient (Fig. 2e). Levels of the other five intermediates downstream of C-22 hydroxylation—6-deoxocathasterone (6-DeoxoCT), 6-deoxoteasterone (6-DeoxoTE), 3-dehydro-6-deoxoteasterone (6-Deoxo3DT), 6-deoxotyphasterol (6-DeoxoTY) and 6-deoxocastasterone (6-DeoxoCS)—were also greatly reduced in osdwarf4-1/d11-4 (Fig. 2e). This indicates that both proteins are involved in C-22 hydroxylation. It is unclear why the upstream compound, campestanol, did not accumulate in osdwarf4-1/d11-4. Because A. thaliana CYP90B1/DWARF4 also catalyzes C-22 hydroxylation of other upstream compounds such as campesterol and (24R)-ergost-4-en-3-one<ref name="ref10" />, it is possible that campestanol is not the major substrate for CYP90B2/OsDWARF4 and CYP724B1/D11 in rice.
[[File:Initpintu_副本.jpg|right|thumb|500px|'''Figure 1.''' ''Phenotypes of rice mutants defective in brassinosteroid biosynthesis.(a) Comparison of gross morphology between the wild-type, single mutants (osdwarf4-1, d11-4, brd1-1), and a double mutant (osdwarf4-1/d11-4). Bar, 30 cm. (b) Erect leaf phenotype of osdwarf4-1. The degree of bending between leaf blade and sheath of osdwarf4-1 (right) is less than that of the wild-type (left). lb, leaf blade; ls, leaf sheath. Bar, 3 cm. (c,d) Close-up views of the osdwarf4-1/d11-4 double mutant (c) and brd1-1 (d). Bars, 5 cm. (e) Endogenous contents of brassinosteroid intermediates in the wild-type and osdwarf4-1/d11-4 double mutant. CN, campestanol; 6-DeoxoCT, 6-deoxocathasterone; 6-DeoxoTE, 6-deoxoteasterone; 6-Deoxo3DT, 3-dehydro-6-deoxoteasterone; 6-DeoxoTY, 6-deoxotyphasterol; 6-DeoxoCS, 6-deoxocastasterone; CS, castasterone; BL, brassinolide.]]
----Characterization of a field-grown osdwarf4-1 mutant

Because the harvest index (grain/(grain plus straw)) of rice is about 0.5, further increases in yield potential will have to involve increases in crop biomass driven by more net photosynthesis<ref name="ref11" />. In rice, the contribution of lower leaves to photosynthesis is significant, even though their photosynthetic capacity is considerably less than that of upper leaves<ref name="ref12" />. Erect leaves allow greater penetration of light to lower leaves, thereby optimizing canopy photosynthesis<ref name="ref13" />. Therefore, the most likely explanation for the increase in above-ground biomass in osdwarf4-1 plots under dense planting conditions (more than 1.2 times that in wild-type under conventional planting condition) is that the shade of the upper leaves is minimized, and the lower leaves receive more light to drive higher rates of photosynthesis (Fig. 3 and Supplementary Table 2 online). No difference was observed in the heading date (flowering time) and the grain-filling period (time to maturity) between wild-type and osdwarf4-1 plants (data not shown). In the wild-type, the above-ground biomass was increased in response to higher fertilizer application under conventional planting (22.2 plants m-2), but was significantly decreased under dense planting with doubled fertilizer application owing to reduced photosynthesis caused by lodging (Fig. 3a). In contrast, the above-ground biomass of osdwarf4-1 was significantly (P < 0.01) increased by dense planting at each fertilizer rate. Consequently, the above-ground biomass in osdwarf4-1 plots under dense planting with standard and a 1.5-fold increase in fertilizer application was approx1.34 and 1.52 times that in wild-type plots under conventional cultivation conditions. The increased biomass depended mainly on accelerated formation of tillers, which have the ability to generate a panicle (panicle number, Fig. 3b). Both total and fertile grain numbers tended to increase in densely planted osdwarf4-1 plots (Fig. 3c), resulting in significantly increased grain yield even when the fertilizer application was not increased (for example, the osdwarf4-1 yield was 6.19 t ha-1 under dense planting with standard fertilizer application whereas the wild-type yield was 4.69 t ha-1 under conventional conditions; Fig. 3d). However, the increase in grain yield was not as much as that in above-ground biomass, because the harvest index of osdwarf4-1 (0.41) was slightly lower than that of the wild-type (0.44). A similar field test gave comparable results.

----Overexpression of OsDWARF4

Overexpression of DWARF in transgenic tomato increases plant height<ref name="ref14" />, and overexpression of DWARF4 in transgenic A. thaliana and tobacco increases vegetative growth and seed yield<ref name="ref15" />. In rice, however, the role of brassinosteroid in regulating leaf angle has been known for over 40 years<ref name="ref16" />,<ref name="ref17" />. Whereas brassinosteroid deficiency increased the erectness of leaves, the opposite phenotype (increased leaf angle) was observed in transgenic rice overexpressing OsDWARF4 (27.6° versus 10.0° in the wild-type, P < 0.01), and was inherited in their progeny (data not shown). Because inclined leaves are unfavorable for photosynthesis when plants are grown in a group, we have to carefully evaluate whether the overexpression of a brassinosteroid-biosynthesis gene in transgenic rice increases grain yield as is the case in transgenic plants overexpressing DWARF4.

===Conclusion===

More erect leaves improve not only light distribution but also the size and use of leaf nitrogen reservoirs for grain growth<ref name="ref13" />. The erect leaf phenotype caused by brassinosteroid deficiency improved biomass production under field conditions, resulting in increased grain yield without the negative environmental effects associated with fertilizers. Recently identified rice homologs of A. thaliana BAS1 that encodes the brassinosteroid catabolic enzyme, CYP734A1 (ref.<ref name="ref18" />). Overexpression of these rice homologs in transgenic rice successfully induced the brassinosteroid-deficient phenotypes (unpublished results). Although ideally, expression of these genes should be restricted to particular tissues and organs such as leaf laminae, it should be possible to generate erect leaf varieties by the introduction of a single transgene. In conclusion, engineering more erect leaves modulating brassinosteroid metabolism can be combined with dense planting to improve rice grain yields.

==Labs working on this gene==
Please input related labs here.

==References==
<references>
<ref name="ref1">Sakamoto, T. & Matsuoka, M. Generating high-yielding varieties by genetic manipulation of plant architecture. Curr. Opin. Biotech. 15, 144–147 (2004).</ref>
<ref name="ref2">Li, J. & Chory, J. A putative leucine-rich repeat receptor kinase involved in brassinosteroid signal transduction. Cell 90, 929–938 (1997).</ref>
<ref name="ref3">Yamamuro, C. et al. Loss of function of a rice brassinosteroid insensitive1 homolog prevents internode elongation and bending of the lamina joint. Plant Cell 12, 1591–1606 (2000).</ref>

<ref name="ref4">Hong, Z. et al. Loss-of-function of a rice brassinosteroid biosynthetic enzyme, C-6 oxidase, prevents the organized arrangement and polar elongation of cells in the leaves and stem. Plant J. 32, 495–508 (2002).</ref>
<ref name="ref5">Hong, Z. et al. A rice brassinosteroid-deficient mutant, ebisu dwarf (d2), is caused by a loss of function of a new member of cytochrome P450. Plant Cell 15, 2900–2910 (2003).</ref>
<ref name="ref6">Sakamoto T;Morinaka Y;Ohnishi T;Sunohara H;Fujioka S;Ueguchi Tanaka M;Mizutani M;Sakata K;Takatsuto S;Yoshida S;Tanaka H;Kitano H;Matsuoka M Erect leaves caused by brassinosteroid deficiency increase biomass production and grain yield in rice Nature Biotechnology, 2006, 24(1): 105-109.</ref>
<ref name="ref7">Choe, S. et al. The DWF4 gene of Arabidopsis encodes a cytochrome P450 that mediates multiple 22alpha-hydroxylation steps in brassinosteroid biosynthesis. Plant Cell 10, 231–243 (1998).</ref>
<ref name="ref8">Bancos, S. et al. Regulation of transcript levels of the Arabidopsis cytochrome P450 genes involved in brassinosteroid biosynthesis. Plant Physiol. 130, 504–513 (2002).</ref>
<ref name="ref9">Tanabe, S. et al. A novel cytochrome P450 is implicated in brassinosteroid biosynthesis via the characterization of a rice dwarf mutant, dwarf11, with reduced seed length. Plant Cell 17, 776–790 (2005).</ref>
<ref name="ref10">Fujioka, S., Takatsuto, S. & Yoshida, S. An early C-22 oxidation branch in brassinosteroid biosynthetic pathway. Plant Physiol. 130, 930–939 (2002).</ref>
<ref name="ref11">Mann, C.C. Crop scientists seek a new revolution. Science 283, 310–314 (1999).</ref>
<ref name="ref12">Horton, P. Prospects for crop improvement through the genetic manipulation of photosynthesis: morphological and biochemical aspects of light capture. J. Exp. Bot. 51, 475–485 (2000).</ref>
<ref name="ref13">Sinclair, T.R. & Sheehy, J.E. Erect leaves and photosynthesis in rice. Science 283, 1455 (1999).</ref>
<ref name="ref14">Bishop, G.J. et al. The tomato DWARF enzyme catalyses C-6 oxidation in brassinosteroid biosynthesis. Proc. Natl. Acad. Sci. USA 96, 1761–1766 (1999).</ref>
<ref name="ref15">Choe, S. et al. Overexpression of DWARF4 in the brassinosteroid biosynthetic pathway results in increased vegetative growth and seed yield in Arabidopsis. Plant J. 26, 573–582 (2001).</ref>
<ref name="ref16">Maeda, E. Rate of lamina inclination in excised rice leaves. Physiol. Plant. 18, 813–827 (1965).</ref>
<ref name="ref17">Wada, K. et al. Brassinolide and homobrassinolide promotion of lamina inclination of rice seedlings. Plant Cell Physiol. 22, 323–325 (1981).</ref>
<ref name="ref18">Neff, M.M. et al. BAS1: A gene regulating brassinosteroid levels and light responsiveness in Arabidopsis. Proc. Natl. Acad. Sci. USA 96, 15316–15323 (1999).</ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os03g0227700|
Description = Cytochrome P450 family protein|
Version = NM_001055982.2 GI:297600576 GeneID:4332134|
Length = 6529 bp|
Definition = Oryza sativa Japonica Group Os03g0227700, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 3|Chromosome 3]]|
AP = Chromosome 3:6765109..6771637|
CDS = 6765109..6765374,6765543..6765867,6765971..6766123,6768937..6769188,6769337..6769429 ,6770222..6770300,6770398..6770504,6771392..6771637|
GCID = <gbrowseImage1>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</cdnaseq>|
AA = <aaseq>MAAMMASITSELLFFLPFILLALLTFYTTTVAKCHGGHWWRGGT TPAKRKRMNLPPGAAGWPLVGETFGYLRAHPATSVGRFMEQHIARYGKIYRSSLFGER TVVSADAGLNRYILQNEGRLFECSYPRSIGGILGKWSMLVLVGDPHREMRAISLNFLS SVRLRAVLLPEVERHTLLVLRAWPPSSTFSAQHQAKKFTFNLMAKNIMSMDPGEEETE RLRREYITFMKGVVSAPLNLPGTPYWKALKSRAAILGVIERKMEERVEKLSKEDASVE QDDLLGWALKQSNLSKEQILDLLLSLLFAGHETSSMALALAIFFLEGCPKAVQELREE HLGIARRQRLRGECKLSWEDYKEMVFTQCVINETLRLGNVVRFLHRKVIKDVHYKGYD IPSGWKILPVLAAVHLDSSLYEDPQRFNPWRWKSSGSSGGLAQSSSFMPYGGGTRLCA GSELAKLEMAVFLHHLVLNFRWELAEPDQAFVFPFVDFPKGLPIRVHRIAQDDEQE</aaseq>|
DNA = <dnaseqindica>1..266#435..759#863..1015#3829..4080#4229..4321#5114..5192#5290..5396#6284..6529#atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtcggtggtttcattactcggtacacgcacacagctatataagcacgcgcactgctcgcctcgccgtgaagcaagcttagctagtagctaggatatcgatcgatcggtaataataggctgacggcgagctggggggtggcgtgcggttgcgtgcgtgcgtgcatgcaggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaaggtacaacacacatcgtcctccattaagttgagcatgcatatatcaatcagctcgctctagcgatcaatatcaatgaaacgcgcgtttgcgtgcgtgcgtgcagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaaggtacgtactcgtagctactagctactccctccgtttcgaaatgtttgacgccgttgactttttatcacatatttgaccgttcgttttatttaaaaaaattaagtaattattaattattttcctatcatttgattcattgttaaatatatttttatgtaggcatataattttacatatctcacaaaagtttttaaataagacgaacggtcaaacatttgctaaaaagtcaacggtgtcaaatatttcgaaacggaggaagtagctagctaccaccccacccgtgcggattgatcgatcgatcgatgtccctctctgcctacacacaccctgcgccccggccgtacgcgcgcaaagtcgtgcgcgcatgagttttactccttccgtttcaaaaaaaaaacgacctagacaagtaatatgatgtaacctagaacaacgaatctatacaggagtatatccaaattcattgtcctagattacatcatatctttgtctatgttgccttttttaggaggagagagtactttacatatgagttatgaccggccggccaggttcctttcttcccctattccgtatgcatactataagcagctgcatataggcatatactactatacccagcttagctaaagcaaacaacttgcaacgacgatcatgtgcaggtccctgttgacagaggttagagatgtaaaccatttttgttgttttcattatctaaagagaaaacaagttactacgacgatcgagatgtacgcaaggggatcatatttatattggagaggagagagagatgatttgcgacttgtgagtgataagtgaatatatatactatatatttgtgtaggagtgacgatgacttccctgtcaatgttttgtttggtatgcacacggggggaaaagagtttagagaaaggaattggctagagagggccaggtttagtttctttaatttttgtttggtggttttgtgtgtagctagtccgcatgcatgatatatcatatatatgcatggcccatcattagatatatatacatgacggcagtgcacttattatatatatgtaggggcatatacaaggtgtatgtgtttagaattgtggatggtccaggaattgttgcagctagttctatactagcatagtgaattagtgatccatatcgagatcaggtccaatcggtcagctcgcacagcgggtcttttagtacgatttgttcggacaaattatcttgcctttgagataatcaacataggattgaaactgcatactagcaatagaccatagctatttggcaatatagaaagctaaaaaatgttgattgaaaggtgcgtgcatgcatgctagcagtatgtcttttaggccaggtccagtttttttttttttctcaaaggcagttgcttaaggtgaaattctagcagaggtcatatgaatatatttattatcttccaaacatgagccaaaacctatatatagttaactcaaaaaaccatgcattggtacttcttctgagtatgcatcctggggactgatcgatcatgcagattcttttctttctttttccctggggacgtactgatcatcactcatgtgagcataaaatgttgcctactttttactgcatgtgaggtcacactcaaatgatctttattgattgcctcagcatgcatgctgctagctttcacgatctctctccgatacacacaccctgtgctgtgctcaacagcatgcaacctggcaggtggaatttttaatctgttggaaagtaaatgaccaaccttcataatttggcatgattatatgctggggaatctatcatttatcatgattcaaccgctgaccaaactctttgccacacacccgaaccacattaatttcgttaagcggatgattagccagcaaccttcacagtttggttttaattagttaatgaaagggtgcttgtagttttattagctgaagagacactgcagtactcctactgagcgttctccttaattgggaaatagatctctgcagtactatcctttaactttaagtgaataacatgtagctaggtctcacacttatcaggcccacacatcagtgatccaagtcaaatggcaatacaccataggagattcttccactaattggctaacaaaaataaactgtgatcactgatcacagctgctaacactataacacattaatttgcatgtctcatgaaatgccagatgagttggattaattatctatcttatgttcagtgccgaggaatatacaaggaaacatcattggtatagttgtggagttctgagtgcattatctcacccagtacggtcaaattaatcctagtctccgtaaatatggttgtgtgcactcttagttaattagtgtagacccggaaattatactttgtcgagaatgttgttatccgtgtgttaagtttcaagtaccctgcaggcctataactaaacactgccaacatggatgtttaagaatgaattatacatacagcgcacttggaaaagaaaaggggtagctacccagctgaagaacacctctctctcatttgaaggtcgacacaatatgtgtcttttctatatcgaaaggaaacgacattttttctcttcaaaacagaaagctatgacaagagacataatcctagactagtaggcggttaggcaacaccatatctgaatttgataggtgtcccacatattttttttcttttcctgataagaaggaacttttcaccgtgagcttatttttttctgttctatgtcgacattagttccatgatgtttactgtgtatttttactgttcccctgcagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggtacagagaattacagattcgaaattaacatctactgtcactagctcagtgctacgtttaactgtagagcaacaattgcagaaagcgaaggcatcagaaatgacatgggaatcgattagttacactcacactgacgcgtgacatgcaggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtaaggccccccctactttgttccttccgatccctcctccctaaacgctactaatataaatcggtgcaaaggctttgtgctagcgtagctcttgctgttttgtttggtcacttgcgtagagagatagcaacttcccgaaatgtcatgtacgaatggagactgaaagctctgcgaaataattgccgggccacgtactacagtacttacgctccctctcttcagtttggatcgatcgagctaagaaggcgcacgcatgtgtgcattgatcgatagagtcgatctgttctgccttgtaagctagccagctgaggttctttcgccttggaagtttctacagggtgaggcccatcgtcttgcaacgatccaaatgccatgacaagtacctgttgttgtaggcttccagtagcacggcctgcccccaagaaggtatctgaatcttctgcattgctgttcagacggcagagcagaacatgccgaaataccattccagtcagttgcagtatatgtgtgtggtggtatcagataaaggagcagagtaatttgaggccttagaaattcagatgagttgctcgacaattgtcgcctaccgatcgagaataagttgcatgcatgtatgtgatgtaattatgggtcttagtcagcaggatagtttagtcgctgcagtactgtagattgtagtacttgcgttcaaagcagacccaaaacagaacttgcattcaggaaacaaaacatgaaaaaagaaatatacagtacttggcactgatatttcttttgtttgggtccttattgcaggtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggtgagtagctcgatttcactcttcttccctgaatcgacctctcctctatctgacagcagaagctaactgaactgaactgaactgaactgaactgaaggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaaggtcagtcgcagtaggattatcagtgaatctaatctattaatctcggtttaattgctgggattcaaggaaaagaaaatactgggacaagcgcggatgtcgctgttgggcagtgtctgtcaatcataagcgaaggggatcatgtcagaatctgcaaaagcagcacaggctaaacgcgtgcttgcctcgagcagcaatcatgcagagaggcaagctccgcgataggaacatgcatggtgaatcactgaacctgatatgaaagtgaatgctcagctcagctcgctcgctcgcgcgcgcatgtgctttctcatttcgccgtgttgcgtgctagctagtcaatggtgttcccacgaattgttttctcttctcatcacacaaagtcagtggtgaaaaccgaggtgcatgtgcggtaatggaacgggggattaggcggcgacgccgtgatggattatgcattcgtgcgagaaaaatccaggagcgttggtccaaatcgcaagcggatcaaaccgaaccaatcagtggacgattggagccctgcagtccatccatcacatctccattcttatccgctgcgactcacggacattagttgttgaggggtgaacgacgcatgtgtgtatgtatgctcgatcgatcgatcgaacaatggatctgtgtgtggtgtagttgccaacgccgacgagattatctgctcgtctgcatgcagccgcgaaaaagacatgtgtccggattatcctgcaggagattgagcaagataatctgctgcgcaaaaactgcagccaatcgatcattccatgacgcgaaaaaaaaaacaataattactactgctgatacgataacttcgcttcgaattgttgacacacaagtgttcttcttttgttcgattgcagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001055982.2 RefSeq:Os03g0227700]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 3]]
[[Category:Chromosome 3]]

File:Initpintu 副本.jpg

2014-05-27T13:25:49Z

LQ:

Os03g0227700

2014-05-27T13:19:01Z

LQ: /* Expression */

Please input one-sentence summary here.
The gene OsDWARF4 was found in young elongating tissues.
==Annotated Information==

Brassinosteroids influence both plant height and leaf erectness in rice<ref name="ref1" />. Arabidopsis thaliana BRASSINOSTEROID INSENSITIVE1 (bri1) mutants, which are deficient in the brassinosteroid receptor, have a severe dwarf phenotype<ref name="ref2" />. The rice mutants d61-1 and d61-2, which are weak mutant alleles of OsBRI1, a homolog of A. thaliana BRI1, have both a semi-dwarf phenotype and more erect leaves<ref name="ref3" />. In rice, an erect leaf phenotype has also been associated with loss-of-function, brassinosteroid-deficient mutants such as brassinosteroid-deficient dwarf1 (brd1), which is defective in OsDWARF (the homolog of tomato CYP85A1/DWARF<ref name="ref4" />), and ebisu dwarf (d2), which is defective in CYP90D2/D2 (ref.<ref name="ref5" />). However, the grain yield of d61, brd1 and d2 mutants is decreased because of morphological alterations in their reproductive development.

===Function===

New cultivars with very erect leaves, which increase light capture for photosynthesis and nitrogen storage for grain filling, may have increased grain yields.The erect leaf phenotype of a rice brassinosteroid-deficient mutant, osdwarf4-1, is associated with enhanced grain yields under conditions of dense planting, even without extra fertilizer. Molecular and biochemical studies reveal that two different cytochrome P450s, CYP90B2/OsDWARF4 and CYP724B1/D11, function redundantly in C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis. Therefore, despite the central role of brassinosteroids in plant growth and development, mutation of OsDWARF4 alone causes only limited defects in brassinosteroid biosynthesis and plant morphology<ref name="ref6" />.

===Expression===
The affected gene, OsDWARF4, was identified by characterizing the site of insertion of the Tos17 retrotransposon.

----Molecular characterization of OsDWARF4 and OsDWARF4L1/D11

OsDWARF4 is a homolog of A. thaliana DWARF4, which encodes a cytochrome P450, CYP90B1 (Fig. 1a). CYP90B1/DWARF4 catalyzes C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis<ref name="ref7" />. Only one DWARF4 homolog was identified in the rice genome. OsDWARF4 is located on the short arm of chromosome 3 (35 cM). Its predicted open reading frame consists of eight exons, and encodes a protein of 502 amino acids (Fig. 1b,c). OsDWARF4 is most closely related to DWARF4 (66.3% amino-acid identity; Fig. 1a). The structure of OsDWARF4 is similar to that of DWARF4 throughout its length: five domains found in cytochrome P450s—proline rich, A (also referred to as dioxygen binding), B (steroid binding), C and heme binding—are highly conserved (Fig. 1c). In wild-type rice, OsDWARF4 transcripts are most abundant in leaf blades and roots, but are also found in all other plant parts tested (Fig. 1d). Transcript levels were decreased by exogenous brassinolide treatment, and increased in brassinosteroid-insensitive d61-3 and brassinosteroid-deficient brd1-1 (Fig. 1e). These results suggest that OsDWARF4 expression is feedback regulated, as is that of A. thaliana brassinosteroid-biosynthesis genes<ref name="ref8" />.
OsDWARF4L1 was identified as an OsDWARF4-like gene by in silico screening of rice DNA databases. OsDWARF4L1 is located on the long arm of chromosome 4 (70.9 cM). A recent study demonstrated that a rice semi-dwarf and short-grain mutant, d11, has a loss-of-function mutation in the OsDWARF4L1 gene<ref name="ref9" />. OsDWARF4L1/D11 shows the highest homology with A. thaliana CYP724A1, and is closely related to rice OsDWARF4 and A. thaliana DWARF4 (47.4%, 37.7% and 41.7% amino-acid identities, respectively; Fig. 1a). In wild-type rice, OsDWARF4L1/D11 transcripts were most abundant in vegetative shoot apices, leaf sheaths and roots, and found at more moderate levels in other organs (Fig. 1d). They were decreased by exogenous brassinolide treatment, and increased in d61-3 and brd1-1 (Fig. 1e). These results indicate that OsDWARF4L1/D11 expression is regulated by a feedback mechanism, as is that of OsDWARF4.
[[File:Figure_1_-_Molecular_characterization_of_OsDWARF4_and_OsDWARF4L1-D11..jpg|right|thumb|500px|'''Figure 1.''' ''Molecular characterization of OsDWARF4 and OsDWARF4L1/D11 (a) Phylogenetic relationships among brassinosteroid-biosynthetic cytochrome P450s of rice, A. thaliana and tomato. Bar, 0.1 amino-acid substitutions per site. (b) Genomic organization of the OsDWARF4 gene. Boxes represent exons, and lines separating boxes represent introns. The arrowhead indicates the site of Tos17 insertion in osdwarf4-1. (c) Relative positions of the major domains in OsDWARF4. All of the major domains found in the cytochrome P450 superfamily are conserved in OsDWARF4. The arrowhead indicates the site of Tos17 insertion in osdwarf4-1. (d) Expression of OsDWARF4 and OsDWARF4L1/D11 in various organs of wild-type rice. The gene encoding Histone H3 was used as a control. (e) Feedback regulation of OsDWARF4 and OsDWARF4L1/D11 in brassinolide-treated wild-type plants (left) and brassinosteroid-related mutants (right). Values under each panel indicate relative amounts of each transcript. (f) Function of recombinant CYP90B2/OsDWARF4 and CYP724B1/D11 proteins. Campesterol (CR) was incubated with each recombinant protein and analyzed by GC-MS. Inset shows that the mass spectrum of the trimethylsilyl ester of each metabolite coincided with that of the trimethylsilyl ether of synthesized (22S)-22-hydroxycampesterol (22-OHCR).]]
----Phenotypes of rice mutants defective in brassinosteroid biosynthesis

In osdwarf4-1, Tos17 insertion into exon 6 causes a premature termination in domain C (arrowhead in Fig. 1b,c). Because a premature stop codon before the heme-binding domain caused loss-of-function of A. thaliana DWARF4 (ref.<ref name="ref7" />), we consider osdwarf4-1 to be a null allele. Unexpectedly, osdwarf4-1 plants show only weak morphological phenotypes, such as a slightly dwarfed stature and more erect leaves (Fig. 2a,b). Both of these phenotypes were complemented by the introduction of an entire OsDWARF4 gene (data not shown). These results seem inconsistent with the fact that rice d61-3 and brd1-1, which are loss-of-function mutants of OsBRI1 and OsDWARF respectively, display severe dwarfing and have malformed leaves (for example, see brd1-1 in Fig. 2a), and suggest functional redundancy in brassinosteroid biosynthesis in rice.
Because null alleles of d11, such as d11-4, also show a weak phenotype<ref name="ref9" />(Fig. 2a),OsDWARF4L1/D11 and OsDWARF4 may function redundantly. To confirm this possibility, researcher produced double mutants by crossing homozygous osdwarf4-1 and d11-4 plants. Among the F2 population, 34 of 576 plants showed severe dwarfing and malformed leaves with tortuous leaf blades, indistinguishable from the brassinosteroid-deficient brd1-1 phenotype (Fig. 2a,c,d). This ratio fits the theoretical 9:3:3:1 (wild-type:osdwarf4-1:d11-4:osdwarf4-1/d11-4) segregation predicted by mendelian inheritance. PCR and sequence analyses confirmed that these plants were osdwarf4-1/d11-4 double mutants (data not shown).
Endogenous levels of brassinosteroid intermediates in brd1 clearly indicate that OsDWARF catalyzes C-6 oxidation<ref name="ref4" />. However, quantification in d2 could not determine the step(s) catalyzed by CYP90D2/D2, because CYP90D2/D2 functions redundantly with another gene, CYP90D3, and loss-of-function of CYP90D2/D2 have caused only limited defects in brassinosteroid biosynthesis<ref name="ref5" />. Similarly, endogenous levels of brassinosteroid intermediates were obviously changed in the osdwarf4-1/d11-4 double mutant relative to osdwarf4-1 or d11 single mutants. Although brassinolide was not detected in either the wild-type or osdwarf4-1/d11-4, another bioactive brassinosteroid, castasterone (CS), was detected in the wild-type but not in osdwarf4-1/d11-4, confirming that osdwarf4-1/d11-4 is brassinosteroid-deficient (Fig. 2e). Levels of the other five intermediates downstream of C-22 hydroxylation—6-deoxocathasterone (6-DeoxoCT), 6-deoxoteasterone (6-DeoxoTE), 3-dehydro-6-deoxoteasterone (6-Deoxo3DT), 6-deoxotyphasterol (6-DeoxoTY) and 6-deoxocastasterone (6-DeoxoCS)—were also greatly reduced in osdwarf4-1/d11-4 (Fig. 2e). This indicates that both proteins are involved in C-22 hydroxylation. It is unclear why the upstream compound, campestanol, did not accumulate in osdwarf4-1/d11-4. Because A. thaliana CYP90B1/DWARF4 also catalyzes C-22 hydroxylation of other upstream compounds such as campesterol and (24R)-ergost-4-en-3-one<ref name="ref10" />, it is possible that campestanol is not the major substrate for CYP90B2/OsDWARF4 and CYP724B1/D11 in rice.

----Characterization of a field-grown osdwarf4-1 mutant

Because the harvest index (grain/(grain plus straw)) of rice is about 0.5, further increases in yield potential will have to involve increases in crop biomass driven by more net photosynthesis<ref name="ref11" />. In rice, the contribution of lower leaves to photosynthesis is significant, even though their photosynthetic capacity is considerably less than that of upper leaves<ref name="ref12" />. Erect leaves allow greater penetration of light to lower leaves, thereby optimizing canopy photosynthesis<ref name="ref13" />. Therefore, the most likely explanation for the increase in above-ground biomass in osdwarf4-1 plots under dense planting conditions (more than 1.2 times that in wild-type under conventional planting condition) is that the shade of the upper leaves is minimized, and the lower leaves receive more light to drive higher rates of photosynthesis (Fig. 3 and Supplementary Table 2 online). No difference was observed in the heading date (flowering time) and the grain-filling period (time to maturity) between wild-type and osdwarf4-1 plants (data not shown). In the wild-type, the above-ground biomass was increased in response to higher fertilizer application under conventional planting (22.2 plants m-2), but was significantly decreased under dense planting with doubled fertilizer application owing to reduced photosynthesis caused by lodging (Fig. 3a). In contrast, the above-ground biomass of osdwarf4-1 was significantly (P < 0.01) increased by dense planting at each fertilizer rate. Consequently, the above-ground biomass in osdwarf4-1 plots under dense planting with standard and a 1.5-fold increase in fertilizer application was approx1.34 and 1.52 times that in wild-type plots under conventional cultivation conditions. The increased biomass depended mainly on accelerated formation of tillers, which have the ability to generate a panicle (panicle number, Fig. 3b). Both total and fertile grain numbers tended to increase in densely planted osdwarf4-1 plots (Fig. 3c), resulting in significantly increased grain yield even when the fertilizer application was not increased (for example, the osdwarf4-1 yield was 6.19 t ha-1 under dense planting with standard fertilizer application whereas the wild-type yield was 4.69 t ha-1 under conventional conditions; Fig. 3d). However, the increase in grain yield was not as much as that in above-ground biomass, because the harvest index of osdwarf4-1 (0.41) was slightly lower than that of the wild-type (0.44). A similar field test gave comparable results.

----Overexpression of OsDWARF4

Overexpression of DWARF in transgenic tomato increases plant height<ref name="ref14" />, and overexpression of DWARF4 in transgenic A. thaliana and tobacco increases vegetative growth and seed yield<ref name="ref15" />. In rice, however, the role of brassinosteroid in regulating leaf angle has been known for over 40 years<ref name="ref16" />,<ref name="ref17" />. Whereas brassinosteroid deficiency increased the erectness of leaves, the opposite phenotype (increased leaf angle) was observed in transgenic rice overexpressing OsDWARF4 (27.6° versus 10.0° in the wild-type, P < 0.01), and was inherited in their progeny (data not shown). Because inclined leaves are unfavorable for photosynthesis when plants are grown in a group, we have to carefully evaluate whether the overexpression of a brassinosteroid-biosynthesis gene in transgenic rice increases grain yield as is the case in transgenic plants overexpressing DWARF4.

===Conclusion===

More erect leaves improve not only light distribution but also the size and use of leaf nitrogen reservoirs for grain growth<ref name="ref13" />. The erect leaf phenotype caused by brassinosteroid deficiency improved biomass production under field conditions, resulting in increased grain yield without the negative environmental effects associated with fertilizers. Recently identified rice homologs of A. thaliana BAS1 that encodes the brassinosteroid catabolic enzyme, CYP734A1 (ref.<ref name="ref18" />). Overexpression of these rice homologs in transgenic rice successfully induced the brassinosteroid-deficient phenotypes (unpublished results). Although ideally, expression of these genes should be restricted to particular tissues and organs such as leaf laminae, it should be possible to generate erect leaf varieties by the introduction of a single transgene. In conclusion, engineering more erect leaves modulating brassinosteroid metabolism can be combined with dense planting to improve rice grain yields.

==Labs working on this gene==
Please input related labs here.

==References==
<references>
<ref name="ref1">Sakamoto, T. & Matsuoka, M. Generating high-yielding varieties by genetic manipulation of plant architecture. Curr. Opin. Biotech. 15, 144–147 (2004).</ref>
<ref name="ref2">Li, J. & Chory, J. A putative leucine-rich repeat receptor kinase involved in brassinosteroid signal transduction. Cell 90, 929–938 (1997).</ref>
<ref name="ref3">Yamamuro, C. et al. Loss of function of a rice brassinosteroid insensitive1 homolog prevents internode elongation and bending of the lamina joint. Plant Cell 12, 1591–1606 (2000).</ref>

<ref name="ref4">Hong, Z. et al. Loss-of-function of a rice brassinosteroid biosynthetic enzyme, C-6 oxidase, prevents the organized arrangement and polar elongation of cells in the leaves and stem. Plant J. 32, 495–508 (2002).</ref>
<ref name="ref5">Hong, Z. et al. A rice brassinosteroid-deficient mutant, ebisu dwarf (d2), is caused by a loss of function of a new member of cytochrome P450. Plant Cell 15, 2900–2910 (2003).</ref>
<ref name="ref6">Sakamoto T;Morinaka Y;Ohnishi T;Sunohara H;Fujioka S;Ueguchi Tanaka M;Mizutani M;Sakata K;Takatsuto S;Yoshida S;Tanaka H;Kitano H;Matsuoka M Erect leaves caused by brassinosteroid deficiency increase biomass production and grain yield in rice Nature Biotechnology, 2006, 24(1): 105-109.</ref>
<ref name="ref7">Choe, S. et al. The DWF4 gene of Arabidopsis encodes a cytochrome P450 that mediates multiple 22alpha-hydroxylation steps in brassinosteroid biosynthesis. Plant Cell 10, 231–243 (1998).</ref>
<ref name="ref8">Bancos, S. et al. Regulation of transcript levels of the Arabidopsis cytochrome P450 genes involved in brassinosteroid biosynthesis. Plant Physiol. 130, 504–513 (2002).</ref>
<ref name="ref9">Tanabe, S. et al. A novel cytochrome P450 is implicated in brassinosteroid biosynthesis via the characterization of a rice dwarf mutant, dwarf11, with reduced seed length. Plant Cell 17, 776–790 (2005).</ref>
<ref name="ref10">Fujioka, S., Takatsuto, S. & Yoshida, S. An early C-22 oxidation branch in brassinosteroid biosynthetic pathway. Plant Physiol. 130, 930–939 (2002).</ref>
<ref name="ref11">Mann, C.C. Crop scientists seek a new revolution. Science 283, 310–314 (1999).</ref>
<ref name="ref12">Horton, P. Prospects for crop improvement through the genetic manipulation of photosynthesis: morphological and biochemical aspects of light capture. J. Exp. Bot. 51, 475–485 (2000).</ref>
<ref name="ref13">Sinclair, T.R. & Sheehy, J.E. Erect leaves and photosynthesis in rice. Science 283, 1455 (1999).</ref>
<ref name="ref14">Bishop, G.J. et al. The tomato DWARF enzyme catalyses C-6 oxidation in brassinosteroid biosynthesis. Proc. Natl. Acad. Sci. USA 96, 1761–1766 (1999).</ref>
<ref name="ref15">Choe, S. et al. Overexpression of DWARF4 in the brassinosteroid biosynthetic pathway results in increased vegetative growth and seed yield in Arabidopsis. Plant J. 26, 573–582 (2001).</ref>
<ref name="ref16">Maeda, E. Rate of lamina inclination in excised rice leaves. Physiol. Plant. 18, 813–827 (1965).</ref>
<ref name="ref17">Wada, K. et al. Brassinolide and homobrassinolide promotion of lamina inclination of rice seedlings. Plant Cell Physiol. 22, 323–325 (1981).</ref>
<ref name="ref18">Neff, M.M. et al. BAS1: A gene regulating brassinosteroid levels and light responsiveness in Arabidopsis. Proc. Natl. Acad. Sci. USA 96, 15316–15323 (1999).</ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os03g0227700|
Description = Cytochrome P450 family protein|
Version = NM_001055982.2 GI:297600576 GeneID:4332134|
Length = 6529 bp|
Definition = Oryza sativa Japonica Group Os03g0227700, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 3|Chromosome 3]]|
AP = Chromosome 3:6765109..6771637|
CDS = 6765109..6765374,6765543..6765867,6765971..6766123,6768937..6769188,6769337..6769429 ,6770222..6770300,6770398..6770504,6771392..6771637|
GCID = <gbrowseImage1>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</cdnaseq>|
AA = <aaseq>MAAMMASITSELLFFLPFILLALLTFYTTTVAKCHGGHWWRGGT TPAKRKRMNLPPGAAGWPLVGETFGYLRAHPATSVGRFMEQHIARYGKIYRSSLFGER TVVSADAGLNRYILQNEGRLFECSYPRSIGGILGKWSMLVLVGDPHREMRAISLNFLS SVRLRAVLLPEVERHTLLVLRAWPPSSTFSAQHQAKKFTFNLMAKNIMSMDPGEEETE RLRREYITFMKGVVSAPLNLPGTPYWKALKSRAAILGVIERKMEERVEKLSKEDASVE QDDLLGWALKQSNLSKEQILDLLLSLLFAGHETSSMALALAIFFLEGCPKAVQELREE HLGIARRQRLRGECKLSWEDYKEMVFTQCVINETLRLGNVVRFLHRKVIKDVHYKGYD IPSGWKILPVLAAVHLDSSLYEDPQRFNPWRWKSSGSSGGLAQSSSFMPYGGGTRLCA GSELAKLEMAVFLHHLVLNFRWELAEPDQAFVFPFVDFPKGLPIRVHRIAQDDEQE</aaseq>|
DNA = <dnaseqindica>1..266#435..759#863..1015#3829..4080#4229..4321#5114..5192#5290..5396#6284..6529#atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtcggtggtttcattactcggtacacgcacacagctatataagcacgcgcactgctcgcctcgccgtgaagcaagcttagctagtagctaggatatcgatcgatcggtaataataggctgacggcgagctggggggtggcgtgcggttgcgtgcgtgcgtgcatgcaggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaaggtacaacacacatcgtcctccattaagttgagcatgcatatatcaatcagctcgctctagcgatcaatatcaatgaaacgcgcgtttgcgtgcgtgcgtgcagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaaggtacgtactcgtagctactagctactccctccgtttcgaaatgtttgacgccgttgactttttatcacatatttgaccgttcgttttatttaaaaaaattaagtaattattaattattttcctatcatttgattcattgttaaatatatttttatgtaggcatataattttacatatctcacaaaagtttttaaataagacgaacggtcaaacatttgctaaaaagtcaacggtgtcaaatatttcgaaacggaggaagtagctagctaccaccccacccgtgcggattgatcgatcgatcgatgtccctctctgcctacacacaccctgcgccccggccgtacgcgcgcaaagtcgtgcgcgcatgagttttactccttccgtttcaaaaaaaaaacgacctagacaagtaatatgatgtaacctagaacaacgaatctatacaggagtatatccaaattcattgtcctagattacatcatatctttgtctatgttgccttttttaggaggagagagtactttacatatgagttatgaccggccggccaggttcctttcttcccctattccgtatgcatactataagcagctgcatataggcatatactactatacccagcttagctaaagcaaacaacttgcaacgacgatcatgtgcaggtccctgttgacagaggttagagatgtaaaccatttttgttgttttcattatctaaagagaaaacaagttactacgacgatcgagatgtacgcaaggggatcatatttatattggagaggagagagagatgatttgcgacttgtgagtgataagtgaatatatatactatatatttgtgtaggagtgacgatgacttccctgtcaatgttttgtttggtatgcacacggggggaaaagagtttagagaaaggaattggctagagagggccaggtttagtttctttaatttttgtttggtggttttgtgtgtagctagtccgcatgcatgatatatcatatatatgcatggcccatcattagatatatatacatgacggcagtgcacttattatatatatgtaggggcatatacaaggtgtatgtgtttagaattgtggatggtccaggaattgttgcagctagttctatactagcatagtgaattagtgatccatatcgagatcaggtccaatcggtcagctcgcacagcgggtcttttagtacgatttgttcggacaaattatcttgcctttgagataatcaacataggattgaaactgcatactagcaatagaccatagctatttggcaatatagaaagctaaaaaatgttgattgaaaggtgcgtgcatgcatgctagcagtatgtcttttaggccaggtccagtttttttttttttctcaaaggcagttgcttaaggtgaaattctagcagaggtcatatgaatatatttattatcttccaaacatgagccaaaacctatatatagttaactcaaaaaaccatgcattggtacttcttctgagtatgcatcctggggactgatcgatcatgcagattcttttctttctttttccctggggacgtactgatcatcactcatgtgagcataaaatgttgcctactttttactgcatgtgaggtcacactcaaatgatctttattgattgcctcagcatgcatgctgctagctttcacgatctctctccgatacacacaccctgtgctgtgctcaacagcatgcaacctggcaggtggaatttttaatctgttggaaagtaaatgaccaaccttcataatttggcatgattatatgctggggaatctatcatttatcatgattcaaccgctgaccaaactctttgccacacacccgaaccacattaatttcgttaagcggatgattagccagcaaccttcacagtttggttttaattagttaatgaaagggtgcttgtagttttattagctgaagagacactgcagtactcctactgagcgttctccttaattgggaaatagatctctgcagtactatcctttaactttaagtgaataacatgtagctaggtctcacacttatcaggcccacacatcagtgatccaagtcaaatggcaatacaccataggagattcttccactaattggctaacaaaaataaactgtgatcactgatcacagctgctaacactataacacattaatttgcatgtctcatgaaatgccagatgagttggattaattatctatcttatgttcagtgccgaggaatatacaaggaaacatcattggtatagttgtggagttctgagtgcattatctcacccagtacggtcaaattaatcctagtctccgtaaatatggttgtgtgcactcttagttaattagtgtagacccggaaattatactttgtcgagaatgttgttatccgtgtgttaagtttcaagtaccctgcaggcctataactaaacactgccaacatggatgtttaagaatgaattatacatacagcgcacttggaaaagaaaaggggtagctacccagctgaagaacacctctctctcatttgaaggtcgacacaatatgtgtcttttctatatcgaaaggaaacgacattttttctcttcaaaacagaaagctatgacaagagacataatcctagactagtaggcggttaggcaacaccatatctgaatttgataggtgtcccacatattttttttcttttcctgataagaaggaacttttcaccgtgagcttatttttttctgttctatgtcgacattagttccatgatgtttactgtgtatttttactgttcccctgcagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggtacagagaattacagattcgaaattaacatctactgtcactagctcagtgctacgtttaactgtagagcaacaattgcagaaagcgaaggcatcagaaatgacatgggaatcgattagttacactcacactgacgcgtgacatgcaggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtaaggccccccctactttgttccttccgatccctcctccctaaacgctactaatataaatcggtgcaaaggctttgtgctagcgtagctcttgctgttttgtttggtcacttgcgtagagagatagcaacttcccgaaatgtcatgtacgaatggagactgaaagctctgcgaaataattgccgggccacgtactacagtacttacgctccctctcttcagtttggatcgatcgagctaagaaggcgcacgcatgtgtgcattgatcgatagagtcgatctgttctgccttgtaagctagccagctgaggttctttcgccttggaagtttctacagggtgaggcccatcgtcttgcaacgatccaaatgccatgacaagtacctgttgttgtaggcttccagtagcacggcctgcccccaagaaggtatctgaatcttctgcattgctgttcagacggcagagcagaacatgccgaaataccattccagtcagttgcagtatatgtgtgtggtggtatcagataaaggagcagagtaatttgaggccttagaaattcagatgagttgctcgacaattgtcgcctaccgatcgagaataagttgcatgcatgtatgtgatgtaattatgggtcttagtcagcaggatagtttagtcgctgcagtactgtagattgtagtacttgcgttcaaagcagacccaaaacagaacttgcattcaggaaacaaaacatgaaaaaagaaatatacagtacttggcactgatatttcttttgtttgggtccttattgcaggtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggtgagtagctcgatttcactcttcttccctgaatcgacctctcctctatctgacagcagaagctaactgaactgaactgaactgaactgaactgaaggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaaggtcagtcgcagtaggattatcagtgaatctaatctattaatctcggtttaattgctgggattcaaggaaaagaaaatactgggacaagcgcggatgtcgctgttgggcagtgtctgtcaatcataagcgaaggggatcatgtcagaatctgcaaaagcagcacaggctaaacgcgtgcttgcctcgagcagcaatcatgcagagaggcaagctccgcgataggaacatgcatggtgaatcactgaacctgatatgaaagtgaatgctcagctcagctcgctcgctcgcgcgcgcatgtgctttctcatttcgccgtgttgcgtgctagctagtcaatggtgttcccacgaattgttttctcttctcatcacacaaagtcagtggtgaaaaccgaggtgcatgtgcggtaatggaacgggggattaggcggcgacgccgtgatggattatgcattcgtgcgagaaaaatccaggagcgttggtccaaatcgcaagcggatcaaaccgaaccaatcagtggacgattggagccctgcagtccatccatcacatctccattcttatccgctgcgactcacggacattagttgttgaggggtgaacgacgcatgtgtgtatgtatgctcgatcgatcgatcgaacaatggatctgtgtgtggtgtagttgccaacgccgacgagattatctgctcgtctgcatgcagccgcgaaaaagacatgtgtccggattatcctgcaggagattgagcaagataatctgctgcgcaaaaactgcagccaatcgatcattccatgacgcgaaaaaaaaaacaataattactactgctgatacgataacttcgcttcgaattgttgacacacaagtgttcttcttttgttcgattgcagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001055982.2 RefSeq:Os03g0227700]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 3]]
[[Category:Chromosome 3]]

Os03g0227700

2014-05-27T13:16:54Z

LQ: /* Expression */

Please input one-sentence summary here.
The gene OsDWARF4 was found in young elongating tissues.
==Annotated Information==

Brassinosteroids influence both plant height and leaf erectness in rice<ref name="ref1" />. Arabidopsis thaliana BRASSINOSTEROID INSENSITIVE1 (bri1) mutants, which are deficient in the brassinosteroid receptor, have a severe dwarf phenotype<ref name="ref2" />. The rice mutants d61-1 and d61-2, which are weak mutant alleles of OsBRI1, a homolog of A. thaliana BRI1, have both a semi-dwarf phenotype and more erect leaves<ref name="ref3" />. In rice, an erect leaf phenotype has also been associated with loss-of-function, brassinosteroid-deficient mutants such as brassinosteroid-deficient dwarf1 (brd1), which is defective in OsDWARF (the homolog of tomato CYP85A1/DWARF<ref name="ref4" />), and ebisu dwarf (d2), which is defective in CYP90D2/D2 (ref.<ref name="ref5" />). However, the grain yield of d61, brd1 and d2 mutants is decreased because of morphological alterations in their reproductive development.

===Function===

New cultivars with very erect leaves, which increase light capture for photosynthesis and nitrogen storage for grain filling, may have increased grain yields.The erect leaf phenotype of a rice brassinosteroid-deficient mutant, osdwarf4-1, is associated with enhanced grain yields under conditions of dense planting, even without extra fertilizer. Molecular and biochemical studies reveal that two different cytochrome P450s, CYP90B2/OsDWARF4 and CYP724B1/D11, function redundantly in C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis. Therefore, despite the central role of brassinosteroids in plant growth and development, mutation of OsDWARF4 alone causes only limited defects in brassinosteroid biosynthesis and plant morphology<ref name="ref6" />.

===Expression===
The affected gene, OsDWARF4, was identified by characterizing the site of insertion of the Tos17 retrotransposon.

----Molecular characterization of OsDWARF4 and OsDWARF4L1/D11

OsDWARF4 is a homolog of A. thaliana DWARF4, which encodes a cytochrome P450, CYP90B1 (Fig. 1a). CYP90B1/DWARF4 catalyzes C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis<ref name="ref7" />. Only one DWARF4 homolog was identified in the rice genome. OsDWARF4 is located on the short arm of chromosome 3 (35 cM). Its predicted open reading frame consists of eight exons, and encodes a protein of 502 amino acids (Fig. 1b,c). OsDWARF4 is most closely related to DWARF4 (66.3% amino-acid identity; Fig. 1a). The structure of OsDWARF4 is similar to that of DWARF4 throughout its length: five domains found in cytochrome P450s—proline rich, A (also referred to as dioxygen binding), B (steroid binding), C and heme binding—are highly conserved (Fig. 1c). In wild-type rice, OsDWARF4 transcripts are most abundant in leaf blades and roots, but are also found in all other plant parts tested (Fig. 1d). Transcript levels were decreased by exogenous brassinolide treatment, and increased in brassinosteroid-insensitive d61-3 and brassinosteroid-deficient brd1-1 (Fig. 1e). These results suggest that OsDWARF4 expression is feedback regulated, as is that of A. thaliana brassinosteroid-biosynthesis genes<ref name="ref8" />.
OsDWARF4L1 was identified as an OsDWARF4-like gene by in silico screening of rice DNA databases. OsDWARF4L1 is located on the long arm of chromosome 4 (70.9 cM). A recent study demonstrated that a rice semi-dwarf and short-grain mutant, d11, has a loss-of-function mutation in the OsDWARF4L1 gene<ref name="ref9" />. OsDWARF4L1/D11 shows the highest homology with A. thaliana CYP724A1, and is closely related to rice OsDWARF4 and A. thaliana DWARF4 (47.4%, 37.7% and 41.7% amino-acid identities, respectively; Fig. 1a). In wild-type rice, OsDWARF4L1/D11 transcripts were most abundant in vegetative shoot apices, leaf sheaths and roots, and found at more moderate levels in other organs (Fig. 1d). They were decreased by exogenous brassinolide treatment, and increased in d61-3 and brd1-1 (Fig. 1e). These results indicate that OsDWARF4L1/D11 expression is regulated by a feedback mechanism, as is that of OsDWARF4.
[[File:Figure_1_-_Molecular_characterization_of_OsDWARF4_and_OsDWARF4L1-D11..jpg|right|thumb|500px|'''Figure 1.''' ''Molecular characterization of OsDWARF4 and OsDWARF4L1/D11]]
----Phenotypes of rice mutants defective in brassinosteroid biosynthesis

In osdwarf4-1, Tos17 insertion into exon 6 causes a premature termination in domain C (arrowhead in Fig. 1b,c). Because a premature stop codon before the heme-binding domain caused loss-of-function of A. thaliana DWARF4 (ref.<ref name="ref7" />), we consider osdwarf4-1 to be a null allele. Unexpectedly, osdwarf4-1 plants show only weak morphological phenotypes, such as a slightly dwarfed stature and more erect leaves (Fig. 2a,b). Both of these phenotypes were complemented by the introduction of an entire OsDWARF4 gene (data not shown). These results seem inconsistent with the fact that rice d61-3 and brd1-1, which are loss-of-function mutants of OsBRI1 and OsDWARF respectively, display severe dwarfing and have malformed leaves (for example, see brd1-1 in Fig. 2a), and suggest functional redundancy in brassinosteroid biosynthesis in rice.
Because null alleles of d11, such as d11-4, also show a weak phenotype<ref name="ref9" />(Fig. 2a),OsDWARF4L1/D11 and OsDWARF4 may function redundantly. To confirm this possibility, researcher produced double mutants by crossing homozygous osdwarf4-1 and d11-4 plants. Among the F2 population, 34 of 576 plants showed severe dwarfing and malformed leaves with tortuous leaf blades, indistinguishable from the brassinosteroid-deficient brd1-1 phenotype (Fig. 2a,c,d). This ratio fits the theoretical 9:3:3:1 (wild-type:osdwarf4-1:d11-4:osdwarf4-1/d11-4) segregation predicted by mendelian inheritance. PCR and sequence analyses confirmed that these plants were osdwarf4-1/d11-4 double mutants (data not shown).
Endogenous levels of brassinosteroid intermediates in brd1 clearly indicate that OsDWARF catalyzes C-6 oxidation<ref name="ref4" />. However, quantification in d2 could not determine the step(s) catalyzed by CYP90D2/D2, because CYP90D2/D2 functions redundantly with another gene, CYP90D3, and loss-of-function of CYP90D2/D2 have caused only limited defects in brassinosteroid biosynthesis<ref name="ref5" />. Similarly, endogenous levels of brassinosteroid intermediates were obviously changed in the osdwarf4-1/d11-4 double mutant relative to osdwarf4-1 or d11 single mutants. Although brassinolide was not detected in either the wild-type or osdwarf4-1/d11-4, another bioactive brassinosteroid, castasterone (CS), was detected in the wild-type but not in osdwarf4-1/d11-4, confirming that osdwarf4-1/d11-4 is brassinosteroid-deficient (Fig. 2e). Levels of the other five intermediates downstream of C-22 hydroxylation—6-deoxocathasterone (6-DeoxoCT), 6-deoxoteasterone (6-DeoxoTE), 3-dehydro-6-deoxoteasterone (6-Deoxo3DT), 6-deoxotyphasterol (6-DeoxoTY) and 6-deoxocastasterone (6-DeoxoCS)—were also greatly reduced in osdwarf4-1/d11-4 (Fig. 2e). This indicates that both proteins are involved in C-22 hydroxylation. It is unclear why the upstream compound, campestanol, did not accumulate in osdwarf4-1/d11-4. Because A. thaliana CYP90B1/DWARF4 also catalyzes C-22 hydroxylation of other upstream compounds such as campesterol and (24R)-ergost-4-en-3-one<ref name="ref10" />, it is possible that campestanol is not the major substrate for CYP90B2/OsDWARF4 and CYP724B1/D11 in rice.

----Characterization of a field-grown osdwarf4-1 mutant

Because the harvest index (grain/(grain plus straw)) of rice is about 0.5, further increases in yield potential will have to involve increases in crop biomass driven by more net photosynthesis<ref name="ref11" />. In rice, the contribution of lower leaves to photosynthesis is significant, even though their photosynthetic capacity is considerably less than that of upper leaves<ref name="ref12" />. Erect leaves allow greater penetration of light to lower leaves, thereby optimizing canopy photosynthesis<ref name="ref13" />. Therefore, the most likely explanation for the increase in above-ground biomass in osdwarf4-1 plots under dense planting conditions (more than 1.2 times that in wild-type under conventional planting condition) is that the shade of the upper leaves is minimized, and the lower leaves receive more light to drive higher rates of photosynthesis (Fig. 3 and Supplementary Table 2 online). No difference was observed in the heading date (flowering time) and the grain-filling period (time to maturity) between wild-type and osdwarf4-1 plants (data not shown). In the wild-type, the above-ground biomass was increased in response to higher fertilizer application under conventional planting (22.2 plants m-2), but was significantly decreased under dense planting with doubled fertilizer application owing to reduced photosynthesis caused by lodging (Fig. 3a). In contrast, the above-ground biomass of osdwarf4-1 was significantly (P < 0.01) increased by dense planting at each fertilizer rate. Consequently, the above-ground biomass in osdwarf4-1 plots under dense planting with standard and a 1.5-fold increase in fertilizer application was approx1.34 and 1.52 times that in wild-type plots under conventional cultivation conditions. The increased biomass depended mainly on accelerated formation of tillers, which have the ability to generate a panicle (panicle number, Fig. 3b). Both total and fertile grain numbers tended to increase in densely planted osdwarf4-1 plots (Fig. 3c), resulting in significantly increased grain yield even when the fertilizer application was not increased (for example, the osdwarf4-1 yield was 6.19 t ha-1 under dense planting with standard fertilizer application whereas the wild-type yield was 4.69 t ha-1 under conventional conditions; Fig. 3d). However, the increase in grain yield was not as much as that in above-ground biomass, because the harvest index of osdwarf4-1 (0.41) was slightly lower than that of the wild-type (0.44). A similar field test gave comparable results.

----Overexpression of OsDWARF4

Overexpression of DWARF in transgenic tomato increases plant height<ref name="ref14" />, and overexpression of DWARF4 in transgenic A. thaliana and tobacco increases vegetative growth and seed yield<ref name="ref15" />. In rice, however, the role of brassinosteroid in regulating leaf angle has been known for over 40 years<ref name="ref16" />,<ref name="ref17" />. Whereas brassinosteroid deficiency increased the erectness of leaves, the opposite phenotype (increased leaf angle) was observed in transgenic rice overexpressing OsDWARF4 (27.6° versus 10.0° in the wild-type, P < 0.01), and was inherited in their progeny (data not shown). Because inclined leaves are unfavorable for photosynthesis when plants are grown in a group, we have to carefully evaluate whether the overexpression of a brassinosteroid-biosynthesis gene in transgenic rice increases grain yield as is the case in transgenic plants overexpressing DWARF4.

===Conclusion===

More erect leaves improve not only light distribution but also the size and use of leaf nitrogen reservoirs for grain growth<ref name="ref13" />. The erect leaf phenotype caused by brassinosteroid deficiency improved biomass production under field conditions, resulting in increased grain yield without the negative environmental effects associated with fertilizers. Recently identified rice homologs of A. thaliana BAS1 that encodes the brassinosteroid catabolic enzyme, CYP734A1 (ref.<ref name="ref18" />). Overexpression of these rice homologs in transgenic rice successfully induced the brassinosteroid-deficient phenotypes (unpublished results). Although ideally, expression of these genes should be restricted to particular tissues and organs such as leaf laminae, it should be possible to generate erect leaf varieties by the introduction of a single transgene. In conclusion, engineering more erect leaves modulating brassinosteroid metabolism can be combined with dense planting to improve rice grain yields.

==Labs working on this gene==
Please input related labs here.

==References==
<references>
<ref name="ref1">Sakamoto, T. & Matsuoka, M. Generating high-yielding varieties by genetic manipulation of plant architecture. Curr. Opin. Biotech. 15, 144–147 (2004).</ref>
<ref name="ref2">Li, J. & Chory, J. A putative leucine-rich repeat receptor kinase involved in brassinosteroid signal transduction. Cell 90, 929–938 (1997).</ref>
<ref name="ref3">Yamamuro, C. et al. Loss of function of a rice brassinosteroid insensitive1 homolog prevents internode elongation and bending of the lamina joint. Plant Cell 12, 1591–1606 (2000).</ref>

<ref name="ref4">Hong, Z. et al. Loss-of-function of a rice brassinosteroid biosynthetic enzyme, C-6 oxidase, prevents the organized arrangement and polar elongation of cells in the leaves and stem. Plant J. 32, 495–508 (2002).</ref>
<ref name="ref5">Hong, Z. et al. A rice brassinosteroid-deficient mutant, ebisu dwarf (d2), is caused by a loss of function of a new member of cytochrome P450. Plant Cell 15, 2900–2910 (2003).</ref>
<ref name="ref6">Sakamoto T;Morinaka Y;Ohnishi T;Sunohara H;Fujioka S;Ueguchi Tanaka M;Mizutani M;Sakata K;Takatsuto S;Yoshida S;Tanaka H;Kitano H;Matsuoka M Erect leaves caused by brassinosteroid deficiency increase biomass production and grain yield in rice Nature Biotechnology, 2006, 24(1): 105-109.</ref>
<ref name="ref7">Choe, S. et al. The DWF4 gene of Arabidopsis encodes a cytochrome P450 that mediates multiple 22alpha-hydroxylation steps in brassinosteroid biosynthesis. Plant Cell 10, 231–243 (1998).</ref>
<ref name="ref8">Bancos, S. et al. Regulation of transcript levels of the Arabidopsis cytochrome P450 genes involved in brassinosteroid biosynthesis. Plant Physiol. 130, 504–513 (2002).</ref>
<ref name="ref9">Tanabe, S. et al. A novel cytochrome P450 is implicated in brassinosteroid biosynthesis via the characterization of a rice dwarf mutant, dwarf11, with reduced seed length. Plant Cell 17, 776–790 (2005).</ref>
<ref name="ref10">Fujioka, S., Takatsuto, S. & Yoshida, S. An early C-22 oxidation branch in brassinosteroid biosynthetic pathway. Plant Physiol. 130, 930–939 (2002).</ref>
<ref name="ref11">Mann, C.C. Crop scientists seek a new revolution. Science 283, 310–314 (1999).</ref>
<ref name="ref12">Horton, P. Prospects for crop improvement through the genetic manipulation of photosynthesis: morphological and biochemical aspects of light capture. J. Exp. Bot. 51, 475–485 (2000).</ref>
<ref name="ref13">Sinclair, T.R. & Sheehy, J.E. Erect leaves and photosynthesis in rice. Science 283, 1455 (1999).</ref>
<ref name="ref14">Bishop, G.J. et al. The tomato DWARF enzyme catalyses C-6 oxidation in brassinosteroid biosynthesis. Proc. Natl. Acad. Sci. USA 96, 1761–1766 (1999).</ref>
<ref name="ref15">Choe, S. et al. Overexpression of DWARF4 in the brassinosteroid biosynthetic pathway results in increased vegetative growth and seed yield in Arabidopsis. Plant J. 26, 573–582 (2001).</ref>
<ref name="ref16">Maeda, E. Rate of lamina inclination in excised rice leaves. Physiol. Plant. 18, 813–827 (1965).</ref>
<ref name="ref17">Wada, K. et al. Brassinolide and homobrassinolide promotion of lamina inclination of rice seedlings. Plant Cell Physiol. 22, 323–325 (1981).</ref>
<ref name="ref18">Neff, M.M. et al. BAS1: A gene regulating brassinosteroid levels and light responsiveness in Arabidopsis. Proc. Natl. Acad. Sci. USA 96, 15316–15323 (1999).</ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os03g0227700|
Description = Cytochrome P450 family protein|
Version = NM_001055982.2 GI:297600576 GeneID:4332134|
Length = 6529 bp|
Definition = Oryza sativa Japonica Group Os03g0227700, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 3|Chromosome 3]]|
AP = Chromosome 3:6765109..6771637|
CDS = 6765109..6765374,6765543..6765867,6765971..6766123,6768937..6769188,6769337..6769429 ,6770222..6770300,6770398..6770504,6771392..6771637|
GCID = <gbrowseImage1>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</cdnaseq>|
AA = <aaseq>MAAMMASITSELLFFLPFILLALLTFYTTTVAKCHGGHWWRGGT TPAKRKRMNLPPGAAGWPLVGETFGYLRAHPATSVGRFMEQHIARYGKIYRSSLFGER TVVSADAGLNRYILQNEGRLFECSYPRSIGGILGKWSMLVLVGDPHREMRAISLNFLS SVRLRAVLLPEVERHTLLVLRAWPPSSTFSAQHQAKKFTFNLMAKNIMSMDPGEEETE RLRREYITFMKGVVSAPLNLPGTPYWKALKSRAAILGVIERKMEERVEKLSKEDASVE QDDLLGWALKQSNLSKEQILDLLLSLLFAGHETSSMALALAIFFLEGCPKAVQELREE HLGIARRQRLRGECKLSWEDYKEMVFTQCVINETLRLGNVVRFLHRKVIKDVHYKGYD IPSGWKILPVLAAVHLDSSLYEDPQRFNPWRWKSSGSSGGLAQSSSFMPYGGGTRLCA GSELAKLEMAVFLHHLVLNFRWELAEPDQAFVFPFVDFPKGLPIRVHRIAQDDEQE</aaseq>|
DNA = <dnaseqindica>1..266#435..759#863..1015#3829..4080#4229..4321#5114..5192#5290..5396#6284..6529#atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtcggtggtttcattactcggtacacgcacacagctatataagcacgcgcactgctcgcctcgccgtgaagcaagcttagctagtagctaggatatcgatcgatcggtaataataggctgacggcgagctggggggtggcgtgcggttgcgtgcgtgcgtgcatgcaggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaaggtacaacacacatcgtcctccattaagttgagcatgcatatatcaatcagctcgctctagcgatcaatatcaatgaaacgcgcgtttgcgtgcgtgcgtgcagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaaggtacgtactcgtagctactagctactccctccgtttcgaaatgtttgacgccgttgactttttatcacatatttgaccgttcgttttatttaaaaaaattaagtaattattaattattttcctatcatttgattcattgttaaatatatttttatgtaggcatataattttacatatctcacaaaagtttttaaataagacgaacggtcaaacatttgctaaaaagtcaacggtgtcaaatatttcgaaacggaggaagtagctagctaccaccccacccgtgcggattgatcgatcgatcgatgtccctctctgcctacacacaccctgcgccccggccgtacgcgcgcaaagtcgtgcgcgcatgagttttactccttccgtttcaaaaaaaaaacgacctagacaagtaatatgatgtaacctagaacaacgaatctatacaggagtatatccaaattcattgtcctagattacatcatatctttgtctatgttgccttttttaggaggagagagtactttacatatgagttatgaccggccggccaggttcctttcttcccctattccgtatgcatactataagcagctgcatataggcatatactactatacccagcttagctaaagcaaacaacttgcaacgacgatcatgtgcaggtccctgttgacagaggttagagatgtaaaccatttttgttgttttcattatctaaagagaaaacaagttactacgacgatcgagatgtacgcaaggggatcatatttatattggagaggagagagagatgatttgcgacttgtgagtgataagtgaatatatatactatatatttgtgtaggagtgacgatgacttccctgtcaatgttttgtttggtatgcacacggggggaaaagagtttagagaaaggaattggctagagagggccaggtttagtttctttaatttttgtttggtggttttgtgtgtagctagtccgcatgcatgatatatcatatatatgcatggcccatcattagatatatatacatgacggcagtgcacttattatatatatgtaggggcatatacaaggtgtatgtgtttagaattgtggatggtccaggaattgttgcagctagttctatactagcatagtgaattagtgatccatatcgagatcaggtccaatcggtcagctcgcacagcgggtcttttagtacgatttgttcggacaaattatcttgcctttgagataatcaacataggattgaaactgcatactagcaatagaccatagctatttggcaatatagaaagctaaaaaatgttgattgaaaggtgcgtgcatgcatgctagcagtatgtcttttaggccaggtccagtttttttttttttctcaaaggcagttgcttaaggtgaaattctagcagaggtcatatgaatatatttattatcttccaaacatgagccaaaacctatatatagttaactcaaaaaaccatgcattggtacttcttctgagtatgcatcctggggactgatcgatcatgcagattcttttctttctttttccctggggacgtactgatcatcactcatgtgagcataaaatgttgcctactttttactgcatgtgaggtcacactcaaatgatctttattgattgcctcagcatgcatgctgctagctttcacgatctctctccgatacacacaccctgtgctgtgctcaacagcatgcaacctggcaggtggaatttttaatctgttggaaagtaaatgaccaaccttcataatttggcatgattatatgctggggaatctatcatttatcatgattcaaccgctgaccaaactctttgccacacacccgaaccacattaatttcgttaagcggatgattagccagcaaccttcacagtttggttttaattagttaatgaaagggtgcttgtagttttattagctgaagagacactgcagtactcctactgagcgttctccttaattgggaaatagatctctgcagtactatcctttaactttaagtgaataacatgtagctaggtctcacacttatcaggcccacacatcagtgatccaagtcaaatggcaatacaccataggagattcttccactaattggctaacaaaaataaactgtgatcactgatcacagctgctaacactataacacattaatttgcatgtctcatgaaatgccagatgagttggattaattatctatcttatgttcagtgccgaggaatatacaaggaaacatcattggtatagttgtggagttctgagtgcattatctcacccagtacggtcaaattaatcctagtctccgtaaatatggttgtgtgcactcttagttaattagtgtagacccggaaattatactttgtcgagaatgttgttatccgtgtgttaagtttcaagtaccctgcaggcctataactaaacactgccaacatggatgtttaagaatgaattatacatacagcgcacttggaaaagaaaaggggtagctacccagctgaagaacacctctctctcatttgaaggtcgacacaatatgtgtcttttctatatcgaaaggaaacgacattttttctcttcaaaacagaaagctatgacaagagacataatcctagactagtaggcggttaggcaacaccatatctgaatttgataggtgtcccacatattttttttcttttcctgataagaaggaacttttcaccgtgagcttatttttttctgttctatgtcgacattagttccatgatgtttactgtgtatttttactgttcccctgcagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggtacagagaattacagattcgaaattaacatctactgtcactagctcagtgctacgtttaactgtagagcaacaattgcagaaagcgaaggcatcagaaatgacatgggaatcgattagttacactcacactgacgcgtgacatgcaggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtaaggccccccctactttgttccttccgatccctcctccctaaacgctactaatataaatcggtgcaaaggctttgtgctagcgtagctcttgctgttttgtttggtcacttgcgtagagagatagcaacttcccgaaatgtcatgtacgaatggagactgaaagctctgcgaaataattgccgggccacgtactacagtacttacgctccctctcttcagtttggatcgatcgagctaagaaggcgcacgcatgtgtgcattgatcgatagagtcgatctgttctgccttgtaagctagccagctgaggttctttcgccttggaagtttctacagggtgaggcccatcgtcttgcaacgatccaaatgccatgacaagtacctgttgttgtaggcttccagtagcacggcctgcccccaagaaggtatctgaatcttctgcattgctgttcagacggcagagcagaacatgccgaaataccattccagtcagttgcagtatatgtgtgtggtggtatcagataaaggagcagagtaatttgaggccttagaaattcagatgagttgctcgacaattgtcgcc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Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001055982.2 RefSeq:Os03g0227700]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 3]]
[[Category:Chromosome 3]]

Os03g0227700

2014-05-27T13:15:09Z

LQ: /* Expression */

Please input one-sentence summary here.
The gene OsDWARF4 was found in young elongating tissues.
==Annotated Information==

Brassinosteroids influence both plant height and leaf erectness in rice<ref name="ref1" />. Arabidopsis thaliana BRASSINOSTEROID INSENSITIVE1 (bri1) mutants, which are deficient in the brassinosteroid receptor, have a severe dwarf phenotype<ref name="ref2" />. The rice mutants d61-1 and d61-2, which are weak mutant alleles of OsBRI1, a homolog of A. thaliana BRI1, have both a semi-dwarf phenotype and more erect leaves<ref name="ref3" />. In rice, an erect leaf phenotype has also been associated with loss-of-function, brassinosteroid-deficient mutants such as brassinosteroid-deficient dwarf1 (brd1), which is defective in OsDWARF (the homolog of tomato CYP85A1/DWARF<ref name="ref4" />), and ebisu dwarf (d2), which is defective in CYP90D2/D2 (ref.<ref name="ref5" />). However, the grain yield of d61, brd1 and d2 mutants is decreased because of morphological alterations in their reproductive development.

===Function===

New cultivars with very erect leaves, which increase light capture for photosynthesis and nitrogen storage for grain filling, may have increased grain yields.The erect leaf phenotype of a rice brassinosteroid-deficient mutant, osdwarf4-1, is associated with enhanced grain yields under conditions of dense planting, even without extra fertilizer. Molecular and biochemical studies reveal that two different cytochrome P450s, CYP90B2/OsDWARF4 and CYP724B1/D11, function redundantly in C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis. Therefore, despite the central role of brassinosteroids in plant growth and development, mutation of OsDWARF4 alone causes only limited defects in brassinosteroid biosynthesis and plant morphology<ref name="ref6" />.

===Expression===
The affected gene, OsDWARF4, was identified by characterizing the site of insertion of the Tos17 retrotransposon.

----Molecular characterization of OsDWARF4 and OsDWARF4L1/D11

OsDWARF4 is a homolog of A. thaliana DWARF4, which encodes a cytochrome P450, CYP90B1 (Fig. 1a). CYP90B1/DWARF4 catalyzes C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis<ref name="ref7" />. Only one DWARF4 homolog was identified in the rice genome. OsDWARF4 is located on the short arm of chromosome 3 (35 cM). Its predicted open reading frame consists of eight exons, and encodes a protein of 502 amino acids (Fig. 1b,c). OsDWARF4 is most closely related to DWARF4 (66.3% amino-acid identity; Fig. 1a). The structure of OsDWARF4 is similar to that of DWARF4 throughout its length: five domains found in cytochrome P450s—proline rich, A (also referred to as dioxygen binding), B (steroid binding), C and heme binding—are highly conserved (Fig. 1c). In wild-type rice, OsDWARF4 transcripts are most abundant in leaf blades and roots, but are also found in all other plant parts tested (Fig. 1d). Transcript levels were decreased by exogenous brassinolide treatment, and increased in brassinosteroid-insensitive d61-3 and brassinosteroid-deficient brd1-1 (Fig. 1e). These results suggest that OsDWARF4 expression is feedback regulated, as is that of A. thaliana brassinosteroid-biosynthesis genes<ref name="ref8" />.
OsDWARF4L1 was identified as an OsDWARF4-like gene by in silico screening of rice DNA databases. OsDWARF4L1 is located on the long arm of chromosome 4 (70.9 cM). A recent study demonstrated that a rice semi-dwarf and short-grain mutant, d11, has a loss-of-function mutation in the OsDWARF4L1 gene<ref name="ref9" />. OsDWARF4L1/D11 shows the highest homology with A. thaliana CYP724A1, and is closely related to rice OsDWARF4 and A. thaliana DWARF4 (47.4%, 37.7% and 41.7% amino-acid identities, respectively; Fig. 1a). In wild-type rice, OsDWARF4L1/D11 transcripts were most abundant in vegetative shoot apices, leaf sheaths and roots, and found at more moderate levels in other organs (Fig. 1d). They were decreased by exogenous brassinolide treatment, and increased in d61-3 and brd1-1 (Fig. 1e). These results indicate that OsDWARF4L1/D11 expression is regulated by a feedback mechanism, as is that of OsDWARF4.
[[File:Figure_1_-_Molecular_characterization_of_OsDWARF4_and_OsDWARF4L1-D11..jpg|right|thumb|500px|'''Figure 1.''']]
----Phenotypes of rice mutants defective in brassinosteroid biosynthesis

In osdwarf4-1, Tos17 insertion into exon 6 causes a premature termination in domain C (arrowhead in Fig. 1b,c). Because a premature stop codon before the heme-binding domain caused loss-of-function of A. thaliana DWARF4 (ref.<ref name="ref7" />), we consider osdwarf4-1 to be a null allele. Unexpectedly, osdwarf4-1 plants show only weak morphological phenotypes, such as a slightly dwarfed stature and more erect leaves (Fig. 2a,b). Both of these phenotypes were complemented by the introduction of an entire OsDWARF4 gene (data not shown). These results seem inconsistent with the fact that rice d61-3 and brd1-1, which are loss-of-function mutants of OsBRI1 and OsDWARF respectively, display severe dwarfing and have malformed leaves (for example, see brd1-1 in Fig. 2a), and suggest functional redundancy in brassinosteroid biosynthesis in rice.
Because null alleles of d11, such as d11-4, also show a weak phenotype<ref name="ref9" />(Fig. 2a),OsDWARF4L1/D11 and OsDWARF4 may function redundantly. To confirm this possibility, researcher produced double mutants by crossing homozygous osdwarf4-1 and d11-4 plants. Among the F2 population, 34 of 576 plants showed severe dwarfing and malformed leaves with tortuous leaf blades, indistinguishable from the brassinosteroid-deficient brd1-1 phenotype (Fig. 2a,c,d). This ratio fits the theoretical 9:3:3:1 (wild-type:osdwarf4-1:d11-4:osdwarf4-1/d11-4) segregation predicted by mendelian inheritance. PCR and sequence analyses confirmed that these plants were osdwarf4-1/d11-4 double mutants (data not shown).
Endogenous levels of brassinosteroid intermediates in brd1 clearly indicate that OsDWARF catalyzes C-6 oxidation<ref name="ref4" />. However, quantification in d2 could not determine the step(s) catalyzed by CYP90D2/D2, because CYP90D2/D2 functions redundantly with another gene, CYP90D3, and loss-of-function of CYP90D2/D2 have caused only limited defects in brassinosteroid biosynthesis<ref name="ref5" />. Similarly, endogenous levels of brassinosteroid intermediates were obviously changed in the osdwarf4-1/d11-4 double mutant relative to osdwarf4-1 or d11 single mutants. Although brassinolide was not detected in either the wild-type or osdwarf4-1/d11-4, another bioactive brassinosteroid, castasterone (CS), was detected in the wild-type but not in osdwarf4-1/d11-4, confirming that osdwarf4-1/d11-4 is brassinosteroid-deficient (Fig. 2e). Levels of the other five intermediates downstream of C-22 hydroxylation—6-deoxocathasterone (6-DeoxoCT), 6-deoxoteasterone (6-DeoxoTE), 3-dehydro-6-deoxoteasterone (6-Deoxo3DT), 6-deoxotyphasterol (6-DeoxoTY) and 6-deoxocastasterone (6-DeoxoCS)—were also greatly reduced in osdwarf4-1/d11-4 (Fig. 2e). This indicates that both proteins are involved in C-22 hydroxylation. It is unclear why the upstream compound, campestanol, did not accumulate in osdwarf4-1/d11-4. Because A. thaliana CYP90B1/DWARF4 also catalyzes C-22 hydroxylation of other upstream compounds such as campesterol and (24R)-ergost-4-en-3-one<ref name="ref10" />, it is possible that campestanol is not the major substrate for CYP90B2/OsDWARF4 and CYP724B1/D11 in rice.

----Characterization of a field-grown osdwarf4-1 mutant

Because the harvest index (grain/(grain plus straw)) of rice is about 0.5, further increases in yield potential will have to involve increases in crop biomass driven by more net photosynthesis<ref name="ref11" />. In rice, the contribution of lower leaves to photosynthesis is significant, even though their photosynthetic capacity is considerably less than that of upper leaves<ref name="ref12" />. Erect leaves allow greater penetration of light to lower leaves, thereby optimizing canopy photosynthesis<ref name="ref13" />. Therefore, the most likely explanation for the increase in above-ground biomass in osdwarf4-1 plots under dense planting conditions (more than 1.2 times that in wild-type under conventional planting condition) is that the shade of the upper leaves is minimized, and the lower leaves receive more light to drive higher rates of photosynthesis (Fig. 3 and Supplementary Table 2 online). No difference was observed in the heading date (flowering time) and the grain-filling period (time to maturity) between wild-type and osdwarf4-1 plants (data not shown). In the wild-type, the above-ground biomass was increased in response to higher fertilizer application under conventional planting (22.2 plants m-2), but was significantly decreased under dense planting with doubled fertilizer application owing to reduced photosynthesis caused by lodging (Fig. 3a). In contrast, the above-ground biomass of osdwarf4-1 was significantly (P < 0.01) increased by dense planting at each fertilizer rate. Consequently, the above-ground biomass in osdwarf4-1 plots under dense planting with standard and a 1.5-fold increase in fertilizer application was approx1.34 and 1.52 times that in wild-type plots under conventional cultivation conditions. The increased biomass depended mainly on accelerated formation of tillers, which have the ability to generate a panicle (panicle number, Fig. 3b). Both total and fertile grain numbers tended to increase in densely planted osdwarf4-1 plots (Fig. 3c), resulting in significantly increased grain yield even when the fertilizer application was not increased (for example, the osdwarf4-1 yield was 6.19 t ha-1 under dense planting with standard fertilizer application whereas the wild-type yield was 4.69 t ha-1 under conventional conditions; Fig. 3d). However, the increase in grain yield was not as much as that in above-ground biomass, because the harvest index of osdwarf4-1 (0.41) was slightly lower than that of the wild-type (0.44). A similar field test gave comparable results.

----Overexpression of OsDWARF4

Overexpression of DWARF in transgenic tomato increases plant height<ref name="ref14" />, and overexpression of DWARF4 in transgenic A. thaliana and tobacco increases vegetative growth and seed yield<ref name="ref15" />. In rice, however, the role of brassinosteroid in regulating leaf angle has been known for over 40 years<ref name="ref16" />,<ref name="ref17" />. Whereas brassinosteroid deficiency increased the erectness of leaves, the opposite phenotype (increased leaf angle) was observed in transgenic rice overexpressing OsDWARF4 (27.6° versus 10.0° in the wild-type, P < 0.01), and was inherited in their progeny (data not shown). Because inclined leaves are unfavorable for photosynthesis when plants are grown in a group, we have to carefully evaluate whether the overexpression of a brassinosteroid-biosynthesis gene in transgenic rice increases grain yield as is the case in transgenic plants overexpressing DWARF4.

===Conclusion===

More erect leaves improve not only light distribution but also the size and use of leaf nitrogen reservoirs for grain growth<ref name="ref13" />. The erect leaf phenotype caused by brassinosteroid deficiency improved biomass production under field conditions, resulting in increased grain yield without the negative environmental effects associated with fertilizers. Recently identified rice homologs of A. thaliana BAS1 that encodes the brassinosteroid catabolic enzyme, CYP734A1 (ref.<ref name="ref18" />). Overexpression of these rice homologs in transgenic rice successfully induced the brassinosteroid-deficient phenotypes (unpublished results). Although ideally, expression of these genes should be restricted to particular tissues and organs such as leaf laminae, it should be possible to generate erect leaf varieties by the introduction of a single transgene. In conclusion, engineering more erect leaves modulating brassinosteroid metabolism can be combined with dense planting to improve rice grain yields.

==Labs working on this gene==
Please input related labs here.

==References==
<references>
<ref name="ref1">Sakamoto, T. & Matsuoka, M. Generating high-yielding varieties by genetic manipulation of plant architecture. Curr. Opin. Biotech. 15, 144–147 (2004).</ref>
<ref name="ref2">Li, J. & Chory, J. A putative leucine-rich repeat receptor kinase involved in brassinosteroid signal transduction. Cell 90, 929–938 (1997).</ref>
<ref name="ref3">Yamamuro, C. et al. Loss of function of a rice brassinosteroid insensitive1 homolog prevents internode elongation and bending of the lamina joint. Plant Cell 12, 1591–1606 (2000).</ref>

<ref name="ref4">Hong, Z. et al. Loss-of-function of a rice brassinosteroid biosynthetic enzyme, C-6 oxidase, prevents the organized arrangement and polar elongation of cells in the leaves and stem. Plant J. 32, 495–508 (2002).</ref>
<ref name="ref5">Hong, Z. et al. A rice brassinosteroid-deficient mutant, ebisu dwarf (d2), is caused by a loss of function of a new member of cytochrome P450. Plant Cell 15, 2900–2910 (2003).</ref>
<ref name="ref6">Sakamoto T;Morinaka Y;Ohnishi T;Sunohara H;Fujioka S;Ueguchi Tanaka M;Mizutani M;Sakata K;Takatsuto S;Yoshida S;Tanaka H;Kitano H;Matsuoka M Erect leaves caused by brassinosteroid deficiency increase biomass production and grain yield in rice Nature Biotechnology, 2006, 24(1): 105-109.</ref>
<ref name="ref7">Choe, S. et al. The DWF4 gene of Arabidopsis encodes a cytochrome P450 that mediates multiple 22alpha-hydroxylation steps in brassinosteroid biosynthesis. Plant Cell 10, 231–243 (1998).</ref>
<ref name="ref8">Bancos, S. et al. Regulation of transcript levels of the Arabidopsis cytochrome P450 genes involved in brassinosteroid biosynthesis. Plant Physiol. 130, 504–513 (2002).</ref>
<ref name="ref9">Tanabe, S. et al. A novel cytochrome P450 is implicated in brassinosteroid biosynthesis via the characterization of a rice dwarf mutant, dwarf11, with reduced seed length. Plant Cell 17, 776–790 (2005).</ref>
<ref name="ref10">Fujioka, S., Takatsuto, S. & Yoshida, S. An early C-22 oxidation branch in brassinosteroid biosynthetic pathway. Plant Physiol. 130, 930–939 (2002).</ref>
<ref name="ref11">Mann, C.C. Crop scientists seek a new revolution. Science 283, 310–314 (1999).</ref>
<ref name="ref12">Horton, P. Prospects for crop improvement through the genetic manipulation of photosynthesis: morphological and biochemical aspects of light capture. J. Exp. Bot. 51, 475–485 (2000).</ref>
<ref name="ref13">Sinclair, T.R. & Sheehy, J.E. Erect leaves and photosynthesis in rice. Science 283, 1455 (1999).</ref>
<ref name="ref14">Bishop, G.J. et al. The tomato DWARF enzyme catalyses C-6 oxidation in brassinosteroid biosynthesis. Proc. Natl. Acad. Sci. USA 96, 1761–1766 (1999).</ref>
<ref name="ref15">Choe, S. et al. Overexpression of DWARF4 in the brassinosteroid biosynthetic pathway results in increased vegetative growth and seed yield in Arabidopsis. Plant J. 26, 573–582 (2001).</ref>
<ref name="ref16">Maeda, E. Rate of lamina inclination in excised rice leaves. Physiol. Plant. 18, 813–827 (1965).</ref>
<ref name="ref17">Wada, K. et al. Brassinolide and homobrassinolide promotion of lamina inclination of rice seedlings. Plant Cell Physiol. 22, 323–325 (1981).</ref>
<ref name="ref18">Neff, M.M. et al. BAS1: A gene regulating brassinosteroid levels and light responsiveness in Arabidopsis. Proc. Natl. Acad. Sci. USA 96, 15316–15323 (1999).</ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os03g0227700|
Description = Cytochrome P450 family protein|
Version = NM_001055982.2 GI:297600576 GeneID:4332134|
Length = 6529 bp|
Definition = Oryza sativa Japonica Group Os03g0227700, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 3|Chromosome 3]]|
AP = Chromosome 3:6765109..6771637|
CDS = 6765109..6765374,6765543..6765867,6765971..6766123,6768937..6769188,6769337..6769429 ,6770222..6770300,6770398..6770504,6771392..6771637|
GCID = <gbrowseImage1>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</cdnaseq>|
AA = <aaseq>MAAMMASITSELLFFLPFILLALLTFYTTTVAKCHGGHWWRGGT TPAKRKRMNLPPGAAGWPLVGETFGYLRAHPATSVGRFMEQHIARYGKIYRSSLFGER TVVSADAGLNRYILQNEGRLFECSYPRSIGGILGKWSMLVLVGDPHREMRAISLNFLS SVRLRAVLLPEVERHTLLVLRAWPPSSTFSAQHQAKKFTFNLMAKNIMSMDPGEEETE RLRREYITFMKGVVSAPLNLPGTPYWKALKSRAAILGVIERKMEERVEKLSKEDASVE QDDLLGWALKQSNLSKEQILDLLLSLLFAGHETSSMALALAIFFLEGCPKAVQELREE HLGIARRQRLRGECKLSWEDYKEMVFTQCVINETLRLGNVVRFLHRKVIKDVHYKGYD IPSGWKILPVLAAVHLDSSLYEDPQRFNPWRWKSSGSSGGLAQSSSFMPYGGGTRLCA GSELAKLEMAVFLHHLVLNFRWELAEPDQAFVFPFVDFPKGLPIRVHRIAQDDEQE</aaseq>|
DNA = <dnaseqindica>1..266#435..759#863..1015#3829..4080#4229..4321#5114..5192#5290..5396#6284..6529#atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtcggtggtttcattactcggtacacgcacacagctatataagcacgcgcactgctcgcctcgccgtgaagcaagcttagctagtagctaggatatcgatcgatcggtaataataggctgacggcgagctggggggtggcgtgcggttgcgtgcgtgcgtgcatgcaggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaaggtacaacacacatcgtcctccattaagttgagcatgcatatatcaatcagctcgctctagcgatcaatatcaatgaaacgcgcgtttgcgtgcgtgcgtgcagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaaggtacgtactcgtagctactagctactccctccgtttcgaaatgtttgacgccgttgactttttatcacatatttgaccgttcgttttatttaaaaaaattaagtaattattaattattttcctatcatttgattcattgttaaatatatttttatgtaggcatataattttacatatctcacaaaagtttttaaataagacgaacggtcaaacatttgctaaaaagtcaacggtgtcaaatatttcgaaacggaggaagtagctagctaccaccccacccgtgcggattgatcgatcgatcgatgtccctctctgcctacacacaccctgcgccccggccgtacgcgcgcaaagtcgtgcgcgcatgagttttactccttccgtttcaaaaaaaaaacgacctagacaagtaatatgatgtaacctagaacaacgaatctatacaggagtatatccaaattcattgtcctagattacatcatatctttgtctatgttgccttttttaggaggagagagtactttacatatgagttatgaccggccggccaggttcctttcttcccctattccgtatgcatactataagcagctgcatataggcatatactactatacccagcttagctaaagcaaacaacttgcaacgacgatcatgtgcaggtccctgttgacagaggttagagatgtaaaccatttttgttgttttcattatctaaagagaaaacaagttactacgacgatcgagatgtacgcaaggggatcatatttatattggagaggagagagagatgatttgcgacttgtgagtgataagtgaatatatatactatatatttgtgtaggagtgacgatgacttccctgtcaatgttttgtttggtatgcacacggggggaaaagagtttagagaaaggaattggctagagagggccaggtttagtttctttaatttttgtttggtggttttgtgtgtagctagtccgcatgcatgatatatcatatatatgcatggcccatcattagatatatatacatgacggcagtgcacttattatatatatgtaggggcatatacaaggtgtatgtgtttagaattgtggatggtccaggaattgttgcagctagttctatactagcatagtgaattagtgatccatatcgagatcaggtccaatcggtcagctcgcacagcgggtcttttagtacgatttgttcggacaaattatcttgcctttgagataatcaacataggattgaaactgcatactagcaatagaccatagctatttggcaatatagaaagctaaaaaatgttgattgaaaggtgcgtgcatgcatgctagcagtatgtcttttaggccaggtccagtttttttttttttctcaaaggcagttgcttaaggtgaaattctagcagaggtcatatgaatatatttattatcttccaaacatgagccaaaacctatatatagttaactcaaaaaaccatgcattggtacttcttctgagtatgcatcctggggactgatcgatcatgcagattcttttctttctttttccctggggacgtactgatcatcactcatgtgagcataaaatgttgcctactttttactgcatgtgaggtcacactcaaatgatctttattgattgcctcagcatgcatgctgctagctttcacgatctctctccgatacacacaccctgtgctgtgctcaacagcatgcaacctggcaggtggaatttttaatctgttggaaagtaaatgaccaaccttcataatttggcatgattatatgctggggaatctatcatttatcatgattcaaccgctgaccaaactctttgccacacacccgaaccacattaatttcgttaagcggatgattagccagcaaccttcacagtttggttttaattagttaatgaaagggtgcttgtagttttattagctgaagagacactgcagtactcctactgagcgttctccttaattgggaaatagatctctgcagtactatcctttaactttaagtgaataacatgtagctaggtctcacacttatcaggcccacacatcagtgatccaagtcaaatggcaatacaccataggagattcttccactaattggctaacaaaaataaactgtgatcactgatcacagctgctaacactataacacattaatttgcatgtctcatgaaatgccagatgagttggattaattatctatcttatgttcagtgccgaggaatatacaaggaaacatcattggtatagttgtggagttctgagtgcattatctcacccagtacggtcaaattaatcctagtctccgtaaatatggttgtgtgcactcttagttaattagtgtagacccggaaattatactttgtcgagaatgttgttatccgtgtgttaagtttcaagtaccctgcaggcctataactaaacactgccaacatggatgtttaagaatgaattatacatacagcgcacttggaaaagaaaaggggtagctacccagctgaagaacacctctctctcatttgaaggtcgacacaatatgtgtcttttctatatcgaaaggaaacgacattttttctcttcaaaacagaaagctatgacaagagacataatcctagactagtaggcggttaggcaacaccatatctgaatttgataggtgtcccacatattttttttcttttcctgataagaaggaacttttcaccgtgagcttatttttttctgttctatgtcgacattagttccatgatgtttactgtgtatttttactgttcccctgcagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggtacagagaattacagattcgaaattaacatctactgtcactagctcagtgctacgtttaactgtagagcaacaattgcagaaagcgaaggcatcagaaatgacatgggaatcgattagttacactcacactgacgcgtgacatgcaggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtaaggccccccctactttgttccttccgatccctcctccctaaacgctactaatataaatcggtgcaaaggctttgtgctagcgtagctcttgctgttttgtttggtcacttgcgtagagagatagcaacttcccgaaatgtcatgtacgaatggagactgaaagctctgcgaaataattgccgggccacgtactacagtacttacgctccctctcttcagtttggatcgatcgagctaagaaggcgcacgcatgtgtgcattgatcgatagagtcgatctgttctgccttgtaagctagccagctgaggttctttcgccttggaagtttctacagggtgaggcccatcgtcttgcaacgatccaaatgccatgacaagtacctgttgttgtaggcttccagtagcacggcctgcccccaagaaggtatctgaatcttctgcattgctgttcagacggcagagcagaacatgccgaaataccattccagtcagttgcagtatatgtgtgtggtggtatcagataaaggagcagagtaatttgaggccttagaaattcagatgagttgctcgacaattgtcgcc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Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001055982.2 RefSeq:Os03g0227700]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 3]]
[[Category:Chromosome 3]]

File:Figure 1 - Molecular characterization of OsDWARF4 and OsDWARF4L1-D11..jpg

2014-05-27T13:13:08Z

LQ: uploaded a new version of "File:Figure 1 - Molecular characterization of OsDWARF4 and OsDWARF4L1-D11..jpg"

Os03g0227700

2014-05-27T12:52:45Z

LQ: /* Annotated Information */

Please input one-sentence summary here.
The gene OsDWARF4 was found in young elongating tissues.
==Annotated Information==

Brassinosteroids influence both plant height and leaf erectness in rice<ref name="ref1" />. Arabidopsis thaliana BRASSINOSTEROID INSENSITIVE1 (bri1) mutants, which are deficient in the brassinosteroid receptor, have a severe dwarf phenotype<ref name="ref2" />. The rice mutants d61-1 and d61-2, which are weak mutant alleles of OsBRI1, a homolog of A. thaliana BRI1, have both a semi-dwarf phenotype and more erect leaves<ref name="ref3" />. In rice, an erect leaf phenotype has also been associated with loss-of-function, brassinosteroid-deficient mutants such as brassinosteroid-deficient dwarf1 (brd1), which is defective in OsDWARF (the homolog of tomato CYP85A1/DWARF<ref name="ref4" />), and ebisu dwarf (d2), which is defective in CYP90D2/D2 (ref.<ref name="ref5" />). However, the grain yield of d61, brd1 and d2 mutants is decreased because of morphological alterations in their reproductive development.

===Function===

New cultivars with very erect leaves, which increase light capture for photosynthesis and nitrogen storage for grain filling, may have increased grain yields.The erect leaf phenotype of a rice brassinosteroid-deficient mutant, osdwarf4-1, is associated with enhanced grain yields under conditions of dense planting, even without extra fertilizer. Molecular and biochemical studies reveal that two different cytochrome P450s, CYP90B2/OsDWARF4 and CYP724B1/D11, function redundantly in C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis. Therefore, despite the central role of brassinosteroids in plant growth and development, mutation of OsDWARF4 alone causes only limited defects in brassinosteroid biosynthesis and plant morphology<ref name="ref6" />.

===Expression===
The affected gene, OsDWARF4, was identified by characterizing the site of insertion of the Tos17 retrotransposon.

----Molecular characterization of OsDWARF4 and OsDWARF4L1/D11

OsDWARF4 is a homolog of A. thaliana DWARF4, which encodes a cytochrome P450, CYP90B1 (Fig. 1a). CYP90B1/DWARF4 catalyzes C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis<ref name="ref7" />. Only one DWARF4 homolog was identified in the rice genome. OsDWARF4 is located on the short arm of chromosome 3 (35 cM). Its predicted open reading frame consists of eight exons, and encodes a protein of 502 amino acids (Fig. 1b,c). OsDWARF4 is most closely related to DWARF4 (66.3% amino-acid identity; Fig. 1a). The structure of OsDWARF4 is similar to that of DWARF4 throughout its length: five domains found in cytochrome P450s—proline rich, A (also referred to as dioxygen binding), B (steroid binding), C and heme binding—are highly conserved (Fig. 1c). In wild-type rice, OsDWARF4 transcripts are most abundant in leaf blades and roots, but are also found in all other plant parts tested (Fig. 1d). Transcript levels were decreased by exogenous brassinolide treatment, and increased in brassinosteroid-insensitive d61-3 and brassinosteroid-deficient brd1-1 (Fig. 1e). These results suggest that OsDWARF4 expression is feedback regulated, as is that of A. thaliana brassinosteroid-biosynthesis genes<ref name="ref8" />.
OsDWARF4L1 was identified as an OsDWARF4-like gene by in silico screening of rice DNA databases. OsDWARF4L1 is located on the long arm of chromosome 4 (70.9 cM). A recent study demonstrated that a rice semi-dwarf and short-grain mutant, d11, has a loss-of-function mutation in the OsDWARF4L1 gene<ref name="ref9" />. OsDWARF4L1/D11 shows the highest homology with A. thaliana CYP724A1, and is closely related to rice OsDWARF4 and A. thaliana DWARF4 (47.4%, 37.7% and 41.7% amino-acid identities, respectively; Fig. 1a). In wild-type rice, OsDWARF4L1/D11 transcripts were most abundant in vegetative shoot apices, leaf sheaths and roots, and found at more moderate levels in other organs (Fig. 1d). They were decreased by exogenous brassinolide treatment, and increased in d61-3 and brd1-1 (Fig. 1e). These results indicate that OsDWARF4L1/D11 expression is regulated by a feedback mechanism, as is that of OsDWARF4.

----Phenotypes of rice mutants defective in brassinosteroid biosynthesis

In osdwarf4-1, Tos17 insertion into exon 6 causes a premature termination in domain C (arrowhead in Fig. 1b,c). Because a premature stop codon before the heme-binding domain caused loss-of-function of A. thaliana DWARF4 (ref.<ref name="ref7" />), we consider osdwarf4-1 to be a null allele. Unexpectedly, osdwarf4-1 plants show only weak morphological phenotypes, such as a slightly dwarfed stature and more erect leaves (Fig. 2a,b). Both of these phenotypes were complemented by the introduction of an entire OsDWARF4 gene (data not shown). These results seem inconsistent with the fact that rice d61-3 and brd1-1, which are loss-of-function mutants of OsBRI1 and OsDWARF respectively, display severe dwarfing and have malformed leaves (for example, see brd1-1 in Fig. 2a), and suggest functional redundancy in brassinosteroid biosynthesis in rice.
Because null alleles of d11, such as d11-4, also show a weak phenotype<ref name="ref9" />(Fig. 2a),OsDWARF4L1/D11 and OsDWARF4 may function redundantly. To confirm this possibility, researcher produced double mutants by crossing homozygous osdwarf4-1 and d11-4 plants. Among the F2 population, 34 of 576 plants showed severe dwarfing and malformed leaves with tortuous leaf blades, indistinguishable from the brassinosteroid-deficient brd1-1 phenotype (Fig. 2a,c,d). This ratio fits the theoretical 9:3:3:1 (wild-type:osdwarf4-1:d11-4:osdwarf4-1/d11-4) segregation predicted by mendelian inheritance. PCR and sequence analyses confirmed that these plants were osdwarf4-1/d11-4 double mutants (data not shown).
Endogenous levels of brassinosteroid intermediates in brd1 clearly indicate that OsDWARF catalyzes C-6 oxidation<ref name="ref4" />. However, quantification in d2 could not determine the step(s) catalyzed by CYP90D2/D2, because CYP90D2/D2 functions redundantly with another gene, CYP90D3, and loss-of-function of CYP90D2/D2 have caused only limited defects in brassinosteroid biosynthesis<ref name="ref5" />. Similarly, endogenous levels of brassinosteroid intermediates were obviously changed in the osdwarf4-1/d11-4 double mutant relative to osdwarf4-1 or d11 single mutants. Although brassinolide was not detected in either the wild-type or osdwarf4-1/d11-4, another bioactive brassinosteroid, castasterone (CS), was detected in the wild-type but not in osdwarf4-1/d11-4, confirming that osdwarf4-1/d11-4 is brassinosteroid-deficient (Fig. 2e). Levels of the other five intermediates downstream of C-22 hydroxylation—6-deoxocathasterone (6-DeoxoCT), 6-deoxoteasterone (6-DeoxoTE), 3-dehydro-6-deoxoteasterone (6-Deoxo3DT), 6-deoxotyphasterol (6-DeoxoTY) and 6-deoxocastasterone (6-DeoxoCS)—were also greatly reduced in osdwarf4-1/d11-4 (Fig. 2e). This indicates that both proteins are involved in C-22 hydroxylation. It is unclear why the upstream compound, campestanol, did not accumulate in osdwarf4-1/d11-4. Because A. thaliana CYP90B1/DWARF4 also catalyzes C-22 hydroxylation of other upstream compounds such as campesterol and (24R)-ergost-4-en-3-one<ref name="ref10" />, it is possible that campestanol is not the major substrate for CYP90B2/OsDWARF4 and CYP724B1/D11 in rice.

----Characterization of a field-grown osdwarf4-1 mutant

Because the harvest index (grain/(grain plus straw)) of rice is about 0.5, further increases in yield potential will have to involve increases in crop biomass driven by more net photosynthesis<ref name="ref11" />. In rice, the contribution of lower leaves to photosynthesis is significant, even though their photosynthetic capacity is considerably less than that of upper leaves<ref name="ref12" />. Erect leaves allow greater penetration of light to lower leaves, thereby optimizing canopy photosynthesis<ref name="ref13" />. Therefore, the most likely explanation for the increase in above-ground biomass in osdwarf4-1 plots under dense planting conditions (more than 1.2 times that in wild-type under conventional planting condition) is that the shade of the upper leaves is minimized, and the lower leaves receive more light to drive higher rates of photosynthesis (Fig. 3 and Supplementary Table 2 online). No difference was observed in the heading date (flowering time) and the grain-filling period (time to maturity) between wild-type and osdwarf4-1 plants (data not shown). In the wild-type, the above-ground biomass was increased in response to higher fertilizer application under conventional planting (22.2 plants m-2), but was significantly decreased under dense planting with doubled fertilizer application owing to reduced photosynthesis caused by lodging (Fig. 3a). In contrast, the above-ground biomass of osdwarf4-1 was significantly (P < 0.01) increased by dense planting at each fertilizer rate. Consequently, the above-ground biomass in osdwarf4-1 plots under dense planting with standard and a 1.5-fold increase in fertilizer application was approx1.34 and 1.52 times that in wild-type plots under conventional cultivation conditions. The increased biomass depended mainly on accelerated formation of tillers, which have the ability to generate a panicle (panicle number, Fig. 3b). Both total and fertile grain numbers tended to increase in densely planted osdwarf4-1 plots (Fig. 3c), resulting in significantly increased grain yield even when the fertilizer application was not increased (for example, the osdwarf4-1 yield was 6.19 t ha-1 under dense planting with standard fertilizer application whereas the wild-type yield was 4.69 t ha-1 under conventional conditions; Fig. 3d). However, the increase in grain yield was not as much as that in above-ground biomass, because the harvest index of osdwarf4-1 (0.41) was slightly lower than that of the wild-type (0.44). A similar field test gave comparable results.

----Overexpression of OsDWARF4

Overexpression of DWARF in transgenic tomato increases plant height<ref name="ref14" />, and overexpression of DWARF4 in transgenic A. thaliana and tobacco increases vegetative growth and seed yield<ref name="ref15" />. In rice, however, the role of brassinosteroid in regulating leaf angle has been known for over 40 years<ref name="ref16" />,<ref name="ref17" />. Whereas brassinosteroid deficiency increased the erectness of leaves, the opposite phenotype (increased leaf angle) was observed in transgenic rice overexpressing OsDWARF4 (27.6° versus 10.0° in the wild-type, P < 0.01), and was inherited in their progeny (data not shown). Because inclined leaves are unfavorable for photosynthesis when plants are grown in a group, we have to carefully evaluate whether the overexpression of a brassinosteroid-biosynthesis gene in transgenic rice increases grain yield as is the case in transgenic plants overexpressing DWARF4.

===Conclusion===

More erect leaves improve not only light distribution but also the size and use of leaf nitrogen reservoirs for grain growth<ref name="ref13" />. The erect leaf phenotype caused by brassinosteroid deficiency improved biomass production under field conditions, resulting in increased grain yield without the negative environmental effects associated with fertilizers. Recently identified rice homologs of A. thaliana BAS1 that encodes the brassinosteroid catabolic enzyme, CYP734A1 (ref.<ref name="ref18" />). Overexpression of these rice homologs in transgenic rice successfully induced the brassinosteroid-deficient phenotypes (unpublished results). Although ideally, expression of these genes should be restricted to particular tissues and organs such as leaf laminae, it should be possible to generate erect leaf varieties by the introduction of a single transgene. In conclusion, engineering more erect leaves modulating brassinosteroid metabolism can be combined with dense planting to improve rice grain yields.

==Labs working on this gene==
Please input related labs here.

==References==
<references>
<ref name="ref1">Sakamoto, T. & Matsuoka, M. Generating high-yielding varieties by genetic manipulation of plant architecture. Curr. Opin. Biotech. 15, 144–147 (2004).</ref>
<ref name="ref2">Li, J. & Chory, J. A putative leucine-rich repeat receptor kinase involved in brassinosteroid signal transduction. Cell 90, 929–938 (1997).</ref>
<ref name="ref3">Yamamuro, C. et al. Loss of function of a rice brassinosteroid insensitive1 homolog prevents internode elongation and bending of the lamina joint. Plant Cell 12, 1591–1606 (2000).</ref>

<ref name="ref4">Hong, Z. et al. Loss-of-function of a rice brassinosteroid biosynthetic enzyme, C-6 oxidase, prevents the organized arrangement and polar elongation of cells in the leaves and stem. Plant J. 32, 495–508 (2002).</ref>
<ref name="ref5">Hong, Z. et al. A rice brassinosteroid-deficient mutant, ebisu dwarf (d2), is caused by a loss of function of a new member of cytochrome P450. Plant Cell 15, 2900–2910 (2003).</ref>
<ref name="ref6">Sakamoto T;Morinaka Y;Ohnishi T;Sunohara H;Fujioka S;Ueguchi Tanaka M;Mizutani M;Sakata K;Takatsuto S;Yoshida S;Tanaka H;Kitano H;Matsuoka M Erect leaves caused by brassinosteroid deficiency increase biomass production and grain yield in rice Nature Biotechnology, 2006, 24(1): 105-109.</ref>
<ref name="ref7">Choe, S. et al. The DWF4 gene of Arabidopsis encodes a cytochrome P450 that mediates multiple 22alpha-hydroxylation steps in brassinosteroid biosynthesis. Plant Cell 10, 231–243 (1998).</ref>
<ref name="ref8">Bancos, S. et al. Regulation of transcript levels of the Arabidopsis cytochrome P450 genes involved in brassinosteroid biosynthesis. Plant Physiol. 130, 504–513 (2002).</ref>
<ref name="ref9">Tanabe, S. et al. A novel cytochrome P450 is implicated in brassinosteroid biosynthesis via the characterization of a rice dwarf mutant, dwarf11, with reduced seed length. Plant Cell 17, 776–790 (2005).</ref>
<ref name="ref10">Fujioka, S., Takatsuto, S. & Yoshida, S. An early C-22 oxidation branch in brassinosteroid biosynthetic pathway. Plant Physiol. 130, 930–939 (2002).</ref>
<ref name="ref11">Mann, C.C. Crop scientists seek a new revolution. Science 283, 310–314 (1999).</ref>
<ref name="ref12">Horton, P. Prospects for crop improvement through the genetic manipulation of photosynthesis: morphological and biochemical aspects of light capture. J. Exp. Bot. 51, 475–485 (2000).</ref>
<ref name="ref13">Sinclair, T.R. & Sheehy, J.E. Erect leaves and photosynthesis in rice. Science 283, 1455 (1999).</ref>
<ref name="ref14">Bishop, G.J. et al. The tomato DWARF enzyme catalyses C-6 oxidation in brassinosteroid biosynthesis. Proc. Natl. Acad. Sci. USA 96, 1761–1766 (1999).</ref>
<ref name="ref15">Choe, S. et al. Overexpression of DWARF4 in the brassinosteroid biosynthetic pathway results in increased vegetative growth and seed yield in Arabidopsis. Plant J. 26, 573–582 (2001).</ref>
<ref name="ref16">Maeda, E. Rate of lamina inclination in excised rice leaves. Physiol. Plant. 18, 813–827 (1965).</ref>
<ref name="ref17">Wada, K. et al. Brassinolide and homobrassinolide promotion of lamina inclination of rice seedlings. Plant Cell Physiol. 22, 323–325 (1981).</ref>
<ref name="ref18">Neff, M.M. et al. BAS1: A gene regulating brassinosteroid levels and light responsiveness in Arabidopsis. Proc. Natl. Acad. Sci. USA 96, 15316–15323 (1999).</ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os03g0227700|
Description = Cytochrome P450 family protein|
Version = NM_001055982.2 GI:297600576 GeneID:4332134|
Length = 6529 bp|
Definition = Oryza sativa Japonica Group Os03g0227700, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 3|Chromosome 3]]|
AP = Chromosome 3:6765109..6771637|
CDS = 6765109..6765374,6765543..6765867,6765971..6766123,6768937..6769188,6769337..6769429 ,6770222..6770300,6770398..6770504,6771392..6771637|
GCID = <gbrowseImage1>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</cdnaseq>|
AA = <aaseq>MAAMMASITSELLFFLPFILLALLTFYTTTVAKCHGGHWWRGGT TPAKRKRMNLPPGAAGWPLVGETFGYLRAHPATSVGRFMEQHIARYGKIYRSSLFGER TVVSADAGLNRYILQNEGRLFECSYPRSIGGILGKWSMLVLVGDPHREMRAISLNFLS SVRLRAVLLPEVERHTLLVLRAWPPSSTFSAQHQAKKFTFNLMAKNIMSMDPGEEETE RLRREYITFMKGVVSAPLNLPGTPYWKALKSRAAILGVIERKMEERVEKLSKEDASVE QDDLLGWALKQSNLSKEQILDLLLSLLFAGHETSSMALALAIFFLEGCPKAVQELREE HLGIARRQRLRGECKLSWEDYKEMVFTQCVINETLRLGNVVRFLHRKVIKDVHYKGYD IPSGWKILPVLAAVHLDSSLYEDPQRFNPWRWKSSGSSGGLAQSSSFMPYGGGTRLCA GSELAKLEMAVFLHHLVLNFRWELAEPDQAFVFPFVDFPKGLPIRVHRIAQDDEQE</aaseq>|
DNA = <dnaseqindica>1..266#435..759#863..1015#3829..4080#4229..4321#5114..5192#5290..5396#6284..6529#atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtcggtggtttcattactcggtacacgcacacagctatataagcacgcgcactgctcgcctcgccgtgaagcaagcttagctagtagctaggatatcgatcgatcggtaataataggctgacggcgagctggggggtggcgtgcggttgcgtgcgtgcgtgcatgcaggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaaggtacaacacacatcgtcctccattaagttgagcatgcatatatcaatcagctcgctctagcgatcaatatcaatgaaacgcgcgtttgcgtgcgtgcgtgcagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaaggtacgtactcgtagctactagctactccctccgtttcgaaatgtttgacgccgttgactttttatcacatatttgaccgttcgttttatttaaaaaaattaagtaattattaattattttcctatcatttgattcattgttaaatatatttttatgtaggcatataattttacatatctcacaaaagtttttaaataagacgaacggtcaaacatttgctaaaaagtcaacggtgtcaaatatttcgaaacggaggaagtagctagctaccaccccacccgtgcggattgatcgatcgatcgatgtccctctctgcctacacacaccctgcgccccggccgtacgcgcgcaaagtcgtgcgcgcatgagttttactccttccgtttcaaaaaaaaaacgacctagacaagtaatatgatgtaacctagaacaacgaatctatacaggagtatatccaaattcattgtcctagattacatcatatctttgtctatgttgccttttttaggaggagagagtactttacatatgagttatgaccggccggccaggttcctttcttcccctattccgtatgcatactataagcagctgcatataggcatatactactatacccagcttagctaaagcaaacaacttgcaacgacgatcatgtgcaggtccctgttgacagaggttagagatgtaaaccatttttgttgttttcattatctaaagagaaaacaagttactacgacgatcgagatgtacgcaaggggatcatatttatattggagaggagagagagatgatttgcgacttgtgagtgataagtgaatatatatactatatatttgtgtaggagtgacgatgacttccctgtcaatgttttgtttggtatgcacacggggggaaaagagtttagagaaaggaattggctagagagggccaggtttagtttctttaatttttgtttggtggttttgtgtgtagctagtccgcatgcatgatatatcatatatatgcatggcccatcattagatatatatacatgacggcagtgcacttattatatatatgtaggggcatatacaaggtgtatgtgtttagaattgtggatggtccaggaattgttgcagctagttctatactagcatagtgaattagtgatccatatcgagatcaggtccaatcggtcagctcgcacagcgggtcttttagtacgatttgttcggacaaattatcttgcctttgagataatcaacataggattgaaactgcatactagcaatagaccatagctatttggcaatatagaaagctaaaaaatgttgattgaaaggtgcgtgcatgcatgctagcagtatgtcttttaggccaggtccagtttttttttttttctcaaaggcagttgcttaaggtgaaattctagcagaggtcatatgaatatatttattatcttccaaacatgagccaaaacctatatatagttaactcaaaaaaccatgcattggtacttcttctgagtatgcatcctggggactgatcgatcatgcagattcttttctttctttttccctggggacgtactgatcatcactcatgtgagcataaaatgttgcctactttttactgcatgtgaggtcacactcaaatgatctttattgattgcctcagcatgcatgctgctagctttcacgatctctctccgatacacacaccctgtgctgtgctcaacagcatgcaacctggcaggtggaatttttaatctgttggaaagtaaatgaccaaccttcataatttggcatgattatatgctggggaatctatcatttatcatgattcaaccgctgaccaaactctttgccacacacccgaaccacattaatttcgttaagcggatgattagccagcaaccttcacagtttggttttaattagttaatgaaagggtgcttgtagttttattagctgaagagacactgcagtactcctactgagcgttctccttaattgggaaatagatctctgcagtactatcctttaactttaagtgaataacatgtagctaggtctcacacttatcaggcccacacatcagtgatccaagtcaaatggcaatacaccataggagattcttccactaattggctaacaaaaataaactgtgatcactgatcacagctgctaacactataacacattaatttgcatgtctcatgaaatgccagatgagttggattaattatctatcttatgttcagtgccgaggaatatacaaggaaacatcattggtatagttgtggagttctgagtgcattatctcacccagtacggtcaaattaatcctagtctccgtaaatatggttgtgtgcactcttagttaattagtgtagacccggaaattatactttgtcgagaatgttgttatccgtgtgttaagtttcaagtaccctgcaggcctataactaaacactgccaacatggatgtttaagaatgaattatacatacagcgcacttggaaaagaaaaggggtagctacccagctgaagaacacctctctctcatttgaaggtcgacacaatatgtgtcttttctatatcgaaaggaaacgacattttttctcttcaaaacagaaagctatgacaagagacataatcctagactagtaggcggttaggcaacaccatatctgaatttgataggtgtcccacatattttttttcttttcctgataagaaggaacttttcaccgtgagcttatttttttctgttctatgtcgacattagttccatgatgtttactgtgtatttttactgttcccctgcagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggtacagagaattacagattcgaaattaacatctactgtcactagctcagtgctacgtttaactgtagagcaacaattgcagaaagcgaaggcatcagaaatgacatgggaatcgattagttacactcacactgacgcgtgacatgcaggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtaaggccccccctactttgttccttccgatccctcctccctaaacgctactaatataaatcggtgcaaaggctttgtgctagcgtagctcttgctgttttgtttggtcacttgcgtagagagatagcaacttcccgaaatgtcatgtacgaatggagactgaaagctctgcgaaataattgccgggccacgtactacagtacttacgctccctctcttcagtttggatcgatcgagctaagaaggcgcacgcatgtgtgcattgatcgatagagtcgatctgttctgccttgtaagctagccagctgaggttctttcgccttggaagtttctacagggtgaggcccatcgtcttgcaacgatccaaatgccatgacaagtacctgttgttgtaggcttccagtagcacggcctgcccccaagaaggtatctgaatcttctgcattgctgttcagacggcagagcagaacatgccgaaataccattccagtcagttgcagtatatgtgtgtggtggtatcagataaaggagcagagtaatttgaggccttagaaattcagatgagttgctcgacaattgtcgcctaccgatcgagaataagttgcatgcatgtatgtgatgtaattatgggtcttagtcagcaggatagtttagtcgctgcagtactgtagattgtagtacttgcgttcaaagcagacccaaaacagaacttgcattcaggaaacaaaacatgaaaaaagaaatatacagtacttggcactgatatttcttttgtttgggtccttattgcaggtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggtgagtagctcgatttcactcttcttccctgaatcgacctctcctctatctgacagcagaagctaactgaactgaactgaactgaactgaactgaaggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaaggtcagtcgcagtaggattatcagtgaatctaatctattaatctcggtttaattgctgggattcaaggaaaagaaaatactgggacaagcgcggatgtcgctgttgggcagtgtctgtcaatcataagcgaaggggatcatgtcagaatctgcaaaagcagcacaggctaaacgcgtgcttgcctcgagcagcaatcatgcagagaggcaagctccgcgataggaacatgcatggtgaatcactgaacctgatatgaaagtgaatgctcagctcagctcgctcgctcgcgcgcgcatgtgctttctcatttcgccgtgttgcgtgctagctagtcaatggtgttcccacgaattgttttctcttctcatcacacaaagtcagtggtgaaaaccgaggtgcatgtgcggtaatggaacgggggattaggcggcgacgccgtgatggattatgcattcgtgcgagaaaaatccaggagcgttggtccaaatcgcaagcggatcaaaccgaaccaatcagtggacgattggagccctgcagtccatccatcacatctccattcttatccgctgcgactcacggacattagttgttgaggggtgaacgacgcatgtgtgtatgtatgctcgatcgatcgatcgaacaatggatctgtgtgtggtgtagttgccaacgccgacgagattatctgctcgtctgcatgcagccgcgaaaaagacatgtgtccggattatcctgcaggagattgagcaagataatctgctgcgcaaaaactgcagccaatcgatcattccatgacgcgaaaaaaaaaacaataattactactgctgatacgataacttcgcttcgaattgttgacacacaagtgttcttcttttgttcgattgcagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001055982.2 RefSeq:Os03g0227700]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 3]]
[[Category:Chromosome 3]]

Os03g0227700

2014-05-27T12:47:02Z

LQ: /* Annotated Information */

Please input one-sentence summary here.
The gene OsDWARF4 was found in young elongating tissues.
==Annotated Information==

Brassinosteroids influence both plant height and leaf erectness in rice<ref name="ref1" />. Arabidopsis thaliana BRASSINOSTEROID INSENSITIVE1 (bri1) mutants, which are deficient in the brassinosteroid receptor, have a severe dwarf phenotype[2]. The rice mutants d61-1 and d61-2, which are weak mutant alleles of OsBRI1, a homolog of A. thaliana BRI1, have both a semi-dwarf phenotype and more erect leaves[3]. In rice, an erect leaf phenotype has also been associated with loss-of-function, brassinosteroid-deficient mutants such as brassinosteroid-deficient dwarf1 (brd1), which is defective in OsDWARF (the homolog of tomato CYP85A1/DWARF[4]), and ebisu dwarf (d2), which is defective in CYP90D2/D2 (ref. [5]). However, the grain yield of d61, brd1 and d2 mutants is decreased because of morphological alterations in their reproductive development.

===Function===

New cultivars with very erect leaves, which increase light capture for photosynthesis and nitrogen storage for grain filling, may have increased grain yields.The erect leaf phenotype of a rice brassinosteroid-deficient mutant, osdwarf4-1, is associated with enhanced grain yields under conditions of dense planting, even without extra fertilizer. Molecular and biochemical studies reveal that two different cytochrome P450s, CYP90B2/OsDWARF4 and CYP724B1/D11, function redundantly in C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis. Therefore, despite the central role of brassinosteroids in plant growth and development, mutation of OsDWARF4 alone causes only limited defects in brassinosteroid biosynthesis and plant morphology[6].

===Expression===
The affected gene, OsDWARF4, was identified by characterizing the site of insertion of the Tos17 retrotransposon.

----Molecular characterization of OsDWARF4 and OsDWARF4L1/D11

OsDWARF4 is a homolog of A. thaliana DWARF4, which encodes a cytochrome P450, CYP90B1 (Fig. 1a). CYP90B1/DWARF4 catalyzes C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis[7]. Only one DWARF4 homolog was identified in the rice genome. OsDWARF4 is located on the short arm of chromosome 3 (35 cM). Its predicted open reading frame consists of eight exons, and encodes a protein of 502 amino acids (Fig. 1b,c). OsDWARF4 is most closely related to DWARF4 (66.3% amino-acid identity; Fig. 1a). The structure of OsDWARF4 is similar to that of DWARF4 throughout its length: five domains found in cytochrome P450s—proline rich, A (also referred to as dioxygen binding), B (steroid binding), C and heme binding—are highly conserved (Fig. 1c). In wild-type rice, OsDWARF4 transcripts are most abundant in leaf blades and roots, but are also found in all other plant parts tested (Fig. 1d). Transcript levels were decreased by exogenous brassinolide treatment, and increased in brassinosteroid-insensitive d61-3 and brassinosteroid-deficient brd1-1 (Fig. 1e). These results suggest that OsDWARF4 expression is feedback regulated, as is that of A. thaliana brassinosteroid-biosynthesis genes[8].
OsDWARF4L1 was identified as an OsDWARF4-like gene by in silico screening of rice DNA databases. OsDWARF4L1 is located on the long arm of chromosome 4 (70.9 cM). A recent study demonstrated that a rice semi-dwarf and short-grain mutant, d11, has a loss-of-function mutation in the OsDWARF4L1 gene[9]. OsDWARF4L1/D11 shows the highest homology with A. thaliana CYP724A1, and is closely related to rice OsDWARF4 and A. thaliana DWARF4 (47.4%, 37.7% and 41.7% amino-acid identities, respectively; Fig. 1a). In wild-type rice, OsDWARF4L1/D11 transcripts were most abundant in vegetative shoot apices, leaf sheaths and roots, and found at more moderate levels in other organs (Fig. 1d). They were decreased by exogenous brassinolide treatment, and increased in d61-3 and brd1-1 (Fig. 1e). These results indicate that OsDWARF4L1/D11 expression is regulated by a feedback mechanism, as is that of OsDWARF4.

----Phenotypes of rice mutants defective in brassinosteroid biosynthesis

In osdwarf4-1, Tos17 insertion into exon 6 causes a premature termination in domain C (arrowhead in Fig. 1b,c). Because a premature stop codon before the heme-binding domain caused loss-of-function of A. thaliana DWARF4 (ref. [7]), we consider osdwarf4-1 to be a null allele. Unexpectedly, osdwarf4-1 plants show only weak morphological phenotypes, such as a slightly dwarfed stature and more erect leaves (Fig. 2a,b). Both of these phenotypes were complemented by the introduction of an entire OsDWARF4 gene (data not shown). These results seem inconsistent with the fact that rice d61-3 and brd1-1, which are loss-of-function mutants of OsBRI1 and OsDWARF respectively, display severe dwarfing and have malformed leaves (for example, see brd1-1 in Fig. 2a), and suggest functional redundancy in brassinosteroid biosynthesis in rice.
Because null alleles of d11, such as d11-4, also show a weak phenotype[9] (Fig. 2a),OsDWARF4L1/D11 and OsDWARF4 may function redundantly. To confirm this possibility, researcher produced double mutants by crossing homozygous osdwarf4-1 and d11-4 plants. Among the F2 population, 34 of 576 plants showed severe dwarfing and malformed leaves with tortuous leaf blades, indistinguishable from the brassinosteroid-deficient brd1-1 phenotype (Fig. 2a,c,d). This ratio fits the theoretical 9:3:3:1 (wild-type:osdwarf4-1:d11-4:osdwarf4-1/d11-4) segregation predicted by mendelian inheritance. PCR and sequence analyses confirmed that these plants were osdwarf4-1/d11-4 double mutants (data not shown).
Endogenous levels of brassinosteroid intermediates in brd1 clearly indicate that OsDWARF catalyzes C-6 oxidation[4]. However, quantification in d2 could not determine the step(s) catalyzed by CYP90D2/D2, because CYP90D2/D2 functions redundantly with another gene, CYP90D3, and loss-of-function of CYP90D2/D2 have caused only limited defects in brassinosteroid biosynthesis[5]. Similarly, endogenous levels of brassinosteroid intermediates were obviously changed in the osdwarf4-1/d11-4 double mutant relative to osdwarf4-1 or d11 single mutants. Although brassinolide was not detected in either the wild-type or osdwarf4-1/d11-4, another bioactive brassinosteroid, castasterone (CS), was detected in the wild-type but not in osdwarf4-1/d11-4, confirming that osdwarf4-1/d11-4 is brassinosteroid-deficient (Fig. 2e). Levels of the other five intermediates downstream of C-22 hydroxylation—6-deoxocathasterone (6-DeoxoCT), 6-deoxoteasterone (6-DeoxoTE), 3-dehydro-6-deoxoteasterone (6-Deoxo3DT), 6-deoxotyphasterol (6-DeoxoTY) and 6-deoxocastasterone (6-DeoxoCS)—were also greatly reduced in osdwarf4-1/d11-4 (Fig. 2e). This indicates that both proteins are involved in C-22 hydroxylation. It is unclear why the upstream compound, campestanol, did not accumulate in osdwarf4-1/d11-4. Because A. thaliana CYP90B1/DWARF4 also catalyzes C-22 hydroxylation of other upstream compounds such as campesterol and (24R)-ergost-4-en-3-one[10], it is possible that campestanol is not the major substrate for CYP90B2/OsDWARF4 and CYP724B1/D11 in rice.

----Characterization of a field-grown osdwarf4-1 mutant

Because the harvest index (grain/(grain plus straw)) of rice is about 0.5, further increases in yield potential will have to involve increases in crop biomass driven by more net photosynthesis[11]. In rice, the contribution of lower leaves to photosynthesis is significant, even though their photosynthetic capacity is considerably less than that of upper leaves[12]. Erect leaves allow greater penetration of light to lower leaves, thereby optimizing canopy photosynthesis[13]. Therefore, the most likely explanation for the increase in above-ground biomass in osdwarf4-1 plots under dense planting conditions (more than 1.2 times that in wild-type under conventional planting condition) is that the shade of the upper leaves is minimized, and the lower leaves receive more light to drive higher rates of photosynthesis (Fig. 3 and Supplementary Table 2 online). No difference was observed in the heading date (flowering time) and the grain-filling period (time to maturity) between wild-type and osdwarf4-1 plants (data not shown). In the wild-type, the above-ground biomass was increased in response to higher fertilizer application under conventional planting (22.2 plants m-2), but was significantly decreased under dense planting with doubled fertilizer application owing to reduced photosynthesis caused by lodging (Fig. 3a). In contrast, the above-ground biomass of osdwarf4-1 was significantly (P < 0.01) increased by dense planting at each fertilizer rate. Consequently, the above-ground biomass in osdwarf4-1 plots under dense planting with standard and a 1.5-fold increase in fertilizer application was approx1.34 and 1.52 times that in wild-type plots under conventional cultivation conditions. The increased biomass depended mainly on accelerated formation of tillers, which have the ability to generate a panicle (panicle number, Fig. 3b). Both total and fertile grain numbers tended to increase in densely planted osdwarf4-1 plots (Fig. 3c), resulting in significantly increased grain yield even when the fertilizer application was not increased (for example, the osdwarf4-1 yield was 6.19 t ha-1 under dense planting with standard fertilizer application whereas the wild-type yield was 4.69 t ha-1 under conventional conditions; Fig. 3d). However, the increase in grain yield was not as much as that in above-ground biomass, because the harvest index of osdwarf4-1 (0.41) was slightly lower than that of the wild-type (0.44). A similar field test gave comparable results.

----Overexpression of OsDWARF4

Overexpression of DWARF in transgenic tomato increases plant height[14], and overexpression of DWARF4 in transgenic A. thaliana and tobacco increases vegetative growth and seed yield[15]. In rice, however, the role of brassinosteroid in regulating leaf angle has been known for over 40 years[16, 17]. Whereas brassinosteroid deficiency increased the erectness of leaves, the opposite phenotype (increased leaf angle) was observed in transgenic rice overexpressing OsDWARF4 (27.6° versus 10.0° in the wild-type, P < 0.01), and was inherited in their progeny (data not shown). Because inclined leaves are unfavorable for photosynthesis when plants are grown in a group, we have to carefully evaluate whether the overexpression of a brassinosteroid-biosynthesis gene in transgenic rice increases grain yield as is the case in transgenic plants overexpressing DWARF4.

===Conclusion===

More erect leaves improve not only light distribution but also the size and use of leaf nitrogen reservoirs for grain growth[13]. The erect leaf phenotype caused by brassinosteroid deficiency improved biomass production under field conditions, resulting in increased grain yield without the negative environmental effects associated with fertilizers. Recently identified rice homologs of A. thaliana BAS1 that encodes the brassinosteroid catabolic enzyme, CYP734A1 (ref. [18]). Overexpression of these rice homologs in transgenic rice successfully induced the brassinosteroid-deficient phenotypes (unpublished results). Although ideally, expression of these genes should be restricted to particular tissues and organs such as leaf laminae, it should be possible to generate erect leaf varieties by the introduction of a single transgene. In conclusion, engineering more erect leaves modulating brassinosteroid metabolism can be combined with dense planting to improve rice grain yields.

==Labs working on this gene==
Please input related labs here.

==References==
<references>
<ref name="ref1">Sakamoto, T. & Matsuoka, M. Generating high-yielding varieties by genetic manipulation of plant architecture. Curr. Opin. Biotech. 15, 144–147 (2004).</ref>
<ref name="ref2">Li, J. & Chory, J. A putative leucine-rich repeat receptor kinase involved in brassinosteroid signal transduction. Cell 90, 929–938 (1997).</ref>
<ref name="ref3">Yamamuro, C. et al. Loss of function of a rice brassinosteroid insensitive1 homolog prevents internode elongation and bending of the lamina joint. Plant Cell 12, 1591–1606 (2000).</ref>

<ref name="ref4">Hong, Z. et al. Loss-of-function of a rice brassinosteroid biosynthetic enzyme, C-6 oxidase, prevents the organized arrangement and polar elongation of cells in the leaves and stem. Plant J. 32, 495–508 (2002).</ref>
<ref name="ref5">Hong, Z. et al. A rice brassinosteroid-deficient mutant, ebisu dwarf (d2), is caused by a loss of function of a new member of cytochrome P450. Plant Cell 15, 2900–2910 (2003).</ref>
<ref name="ref6">Sakamoto T;Morinaka Y;Ohnishi T;Sunohara H;Fujioka S;Ueguchi Tanaka M;Mizutani M;Sakata K;Takatsuto S;Yoshida S;Tanaka H;Kitano H;Matsuoka M Erect leaves caused by brassinosteroid deficiency increase biomass production and grain yield in rice Nature Biotechnology, 2006, 24(1): 105-109.</ref>
<ref name="ref7">Choe, S. et al. The DWF4 gene of Arabidopsis encodes a cytochrome P450 that mediates multiple 22alpha-hydroxylation steps in brassinosteroid biosynthesis. Plant Cell 10, 231–243 (1998).</ref>
<ref name="ref8">Bancos, S. et al. Regulation of transcript levels of the Arabidopsis cytochrome P450 genes involved in brassinosteroid biosynthesis. Plant Physiol. 130, 504–513 (2002).</ref>
<ref name="ref9">Tanabe, S. et al. A novel cytochrome P450 is implicated in brassinosteroid biosynthesis via the characterization of a rice dwarf mutant, dwarf11, with reduced seed length. Plant Cell 17, 776–790 (2005).</ref>
<ref name="ref10">Fujioka, S., Takatsuto, S. & Yoshida, S. An early C-22 oxidation branch in brassinosteroid biosynthetic pathway. Plant Physiol. 130, 930–939 (2002).</ref>
<ref name="ref11">Mann, C.C. Crop scientists seek a new revolution. Science 283, 310–314 (1999).</ref>
<ref name="ref12">Horton, P. Prospects for crop improvement through the genetic manipulation of photosynthesis: morphological and biochemical aspects of light capture. J. Exp. Bot. 51, 475–485 (2000).</ref>
<ref name="ref13">Sinclair, T.R. & Sheehy, J.E. Erect leaves and photosynthesis in rice. Science 283, 1455 (1999).</ref>
<ref name="ref14">Bishop, G.J. et al. The tomato DWARF enzyme catalyses C-6 oxidation in brassinosteroid biosynthesis. Proc. Natl. Acad. Sci. USA 96, 1761–1766 (1999).</ref>
<ref name="ref15">Choe, S. et al. Overexpression of DWARF4 in the brassinosteroid biosynthetic pathway results in increased vegetative growth and seed yield in Arabidopsis. Plant J. 26, 573–582 (2001).</ref>
<ref name="ref16">Maeda, E. Rate of lamina inclination in excised rice leaves. Physiol. Plant. 18, 813–827 (1965).</ref>
<ref name="ref17">Wada, K. et al. Brassinolide and homobrassinolide promotion of lamina inclination of rice seedlings. Plant Cell Physiol. 22, 323–325 (1981).</ref>
<ref name="ref18">Neff, M.M. et al. BAS1: A gene regulating brassinosteroid levels and light responsiveness in Arabidopsis. Proc. Natl. Acad. Sci. USA 96, 15316–15323 (1999).</ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os03g0227700|
Description = Cytochrome P450 family protein|
Version = NM_001055982.2 GI:297600576 GeneID:4332134|
Length = 6529 bp|
Definition = Oryza sativa Japonica Group Os03g0227700, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 3|Chromosome 3]]|
AP = Chromosome 3:6765109..6771637|
CDS = 6765109..6765374,6765543..6765867,6765971..6766123,6768937..6769188,6769337..6769429 ,6770222..6770300,6770398..6770504,6771392..6771637|
GCID = <gbrowseImage1>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</cdnaseq>|
AA = <aaseq>MAAMMASITSELLFFLPFILLALLTFYTTTVAKCHGGHWWRGGT TPAKRKRMNLPPGAAGWPLVGETFGYLRAHPATSVGRFMEQHIARYGKIYRSSLFGER TVVSADAGLNRYILQNEGRLFECSYPRSIGGILGKWSMLVLVGDPHREMRAISLNFLS SVRLRAVLLPEVERHTLLVLRAWPPSSTFSAQHQAKKFTFNLMAKNIMSMDPGEEETE RLRREYITFMKGVVSAPLNLPGTPYWKALKSRAAILGVIERKMEERVEKLSKEDASVE QDDLLGWALKQSNLSKEQILDLLLSLLFAGHETSSMALALAIFFLEGCPKAVQELREE HLGIARRQRLRGECKLSWEDYKEMVFTQCVINETLRLGNVVRFLHRKVIKDVHYKGYD IPSGWKILPVLAAVHLDSSLYEDPQRFNPWRWKSSGSSGGLAQSSSFMPYGGGTRLCA GSELAKLEMAVFLHHLVLNFRWELAEPDQAFVFPFVDFPKGLPIRVHRIAQDDEQE</aaseq>|
DNA = <dnaseqindica>1..266#435..759#863..1015#3829..4080#4229..4321#5114..5192#5290..5396#6284..6529#atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtcggtggtttcattactcggtacacgcacacagctatataagcacgcgcactgctcgcctcgccgtgaagcaagcttagctagtagctaggatatcgatcgatcggtaataataggctgacggcgagctggggggtggcgtgcggttgcgtgcgtgcgtgcatgcaggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaaggtacaacacacatcgtcctccattaagttgagcatgcatatatcaatcagctcgctctagcgatcaatatcaatgaaacgcgcgtttgcgtgcgtgcgtgcagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaaggtacgtactcgtagctactagctactccctccgtttcgaaatgtttgacgccgttgactttttatcacatatttgaccgttcgttttatttaaaaaaattaagtaattattaattattttcctatcatttgattcattgttaaatatatttttatgtaggcatataattttacatatctcacaaaagtttttaaataagacgaacggtcaaacatttgctaaaaagtcaacggtgtcaaatatttcgaaacggaggaagtagctagctaccaccccacccgtgcggattgatcgatcgatcgatgtccctctctgcctacacacaccctgcgccccggccgtacgcgcgcaaagtcgtgcgcgcatgagttttactccttccgtttcaaaaaaaaaacgacctagacaagtaatatgatgtaacctagaacaacgaatctatacaggagtatatccaaattcattgtcctagattacatcatatctttgtctatgttgccttttttaggaggagagagtactttacatatgagttatgaccggccggccaggttcctttcttcccctattccgtatgcatactataagcagctgcatataggcatatactactatacccagcttagctaaagcaaacaacttgcaacgacgatcatgtgcaggtccctgttgacagaggttagagatgtaaaccatttttgttgttttcattatctaaagagaaaacaagttactacgacgatcgagatgtacgcaaggggatcatatttatattggagaggagagagagatgatttgcgacttgtgagtgataagtgaatatatatactatatatttgtgtaggagtgacgatgacttccctgtcaatgttttgtttggtatgcacacggggggaaaagagtttagagaaaggaattggctagagagggccaggtttagtttctttaatttttgtttggtggttttgtgtgtagctagtccgcatgcatgatatatcatatatatgcatggcccatcattagatatatatacatgacggcagtgcacttattatatatatgtaggggcatatacaaggtgtatgtgtttagaattgtggatggtccaggaattgttgcagctagttctatactagcatagtgaattagtgatccatatcgagatcaggtccaatcggtcagctcgcacagcgggtcttttagtacgatttgttcggacaaattatcttgcctttgagataatcaacataggattgaaactgcatactagcaatagaccatagctatttggcaatatagaaagctaaaaaatgttgattgaaaggtgcgtgcatgcatgctagcagtatgtcttttaggccaggtccagtttttttttttttctcaaaggcagttgcttaaggtgaaattctagcagaggtcatatgaatatatttattatcttccaaacatgagccaaaacctatatatagttaactcaaaaaaccatgcattggtacttcttctgagtatgcatcctggggactgatcgatcatgcagattcttttctttctttttccctggggacgtactgatcatcactcatgtgagcataaaatgttgcctactttttactgcatgtgaggtcacactcaaatgatctttattgattgcctcagcatgcatgctgctagctttcacgatctctctccgatacacacaccctgtgctgtgctcaacagcatgcaacctggcaggtggaatttttaatctgttggaaagtaaatgaccaaccttcataatttggcatgattatatgctggggaatctatcatttatcatgattcaaccgctgaccaaactctttgccacacacccgaaccacattaatttcgttaagcggatgattagccagcaaccttcacagtttggttttaattagttaatgaaagggtgcttgtagttttattagctgaagagacactgcagtactcctactgagcgttctccttaattgggaaatagatctctgcagtactatcctttaactttaagtgaataacatgtagctaggtctcacacttatcaggcccacacatcagtgatccaagtcaaatggcaatacaccataggagattcttccactaattggctaacaaaaataaactgtgatcactgatcacagctgctaacactataacacattaatttgcatgtctcatgaaatgccagatgagttggattaattatctatcttatgttcagtgccgaggaatatacaaggaaacatcattggtatagttgtggagttctgagtgcattatctcacccagtacggtcaaattaatcctagtctccgtaaatatggttgtgtgcactcttagttaattagtgtagacccggaaattatactttgtcgagaatgttgttatccgtgtgttaagtttcaagtaccctgcaggcctataactaaacactgccaacatggatgtttaagaatgaattatacatacagcgcacttggaaaagaaaaggggtagctacccagctgaagaacacctctctctcatttgaaggtcgacacaatatgtgtcttttctatatcgaaaggaaacgacattttttctcttcaaaacagaaagctatgacaagagacataatcctagactagtaggcggttaggcaacaccatatctgaatttgataggtgtcccacatattttttttcttttcctgataagaaggaacttttcaccgtgagcttatttttttctgttctatgtcgacattagttccatgatgtttactgtgtatttttactgttcccctgcagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggtacagagaattacagattcgaaattaacatctactgtcactagctcagtgctacgtttaactgtagagcaacaattgcagaaagcgaaggcatcagaaatgacatgggaatcgattagttacactcacactgacgcgtgacatgcaggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtaaggccccccctactttgttccttccgatccctcctccctaaacgctactaatataaatcggtgcaaaggctttgtgctagcgtagctcttgctgttttgtttggtcacttgcgtagagagatagcaacttcccgaaatgtcatgtacgaatggagactgaaagctctgcgaaataattgccgggccacgtactacagtacttacgctccctctcttcagtttggatcgatcgagctaagaaggcgcacgcatgtgtgcattgatcgatagagtcgatctgttctgccttgtaagctagccagctgaggttctttcgccttggaagtttctacagggtgaggcccatcgtcttgcaacgatccaaatgccatgacaagtacctgttgttgtaggcttccagtagcacggcctgcccccaagaaggtatctgaatcttctgcattgctgttcagacggcagagcagaacatgccgaaataccattccagtcagttgcagtatatgtgtgtggtggtatcagataaaggagcagagtaatttgaggccttagaaattcagatgagttgctcgacaattgtcgcctaccgatcgagaataagttgcatgcatgtatgtgatgtaattatgggtcttagtcagcaggatagtttagtcgctgcagtactgtagattgtagtacttgcgttcaaagcagacccaaaacagaacttgcattcaggaaacaaaacatgaaaaaagaaatatacagtacttggcactgatatttcttttgtttgggtccttattgcaggtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggtgagtagctcgatttcactcttcttccctgaatcgacctctcctctatctgacagcagaagctaactgaactgaactgaactgaactgaactgaaggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaaggtcagtcgcagtaggattatcagtgaatctaatctattaatctcggtttaattgctgggattcaaggaaaagaaaatactgggacaagcgcggatgtcgctgttgggcagtgtctgtcaatcataagcgaaggggatcatgtcagaatctgcaaaagcagcacaggctaaacgcgtgcttgcctcgagcagcaatcatgcagagaggcaagctccgcgataggaacatgcatggtgaatcactgaacctgatatgaaagtgaatgctcagctcagctcgctcgctcgcgcgcgcatgtgctttctcatttcgccgtgttgcgtgctagctagtcaatggtgttcccacgaattgttttctcttctcatcacacaaagtcagtggtgaaaaccgaggtgcatgtgcggtaatggaacgggggattaggcggcgacgccgtgatggattatgcattcgtgcgagaaaaatccaggagcgttggtccaaatcgcaagcggatcaaaccgaaccaatcagtggacgattggagccctgcagtccatccatcacatctccattcttatccgctgcgactcacggacattagttgttgaggggtgaacgacgcatgtgtgtatgtatgctcgatcgatcgatcgaacaatggatctgtgtgtggtgtagttgccaacgccgacgagattatctgctcgtctgcatgcagccgcgaaaaagacatgtgtccggattatcctgcaggagattgagcaagataatctgctgcgcaaaaactgcagccaatcgatcattccatgacgcgaaaaaaaaaacaataattactactgctgatacgataacttcgcttcgaattgttgacacacaagtgttcttcttttgttcgattgcagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001055982.2 RefSeq:Os03g0227700]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 3]]
[[Category:Chromosome 3]]

Os03g0227700

2014-05-27T12:45:07Z

LQ: /* References */

Please input one-sentence summary here.
The gene OsDWARF4 was found in young elongating tissues.
==Annotated Information==

Brassinosteroids influence both plant height and leaf erectness in rice[1]. Arabidopsis thaliana BRASSINOSTEROID INSENSITIVE1 (bri1) mutants, which are deficient in the brassinosteroid receptor, have a severe dwarf phenotype[2]. The rice mutants d61-1 and d61-2, which are weak mutant alleles of OsBRI1, a homolog of A. thaliana BRI1, have both a semi-dwarf phenotype and more erect leaves[3]. In rice, an erect leaf phenotype has also been associated with loss-of-function, brassinosteroid-deficient mutants such as brassinosteroid-deficient dwarf1 (brd1), which is defective in OsDWARF (the homolog of tomato CYP85A1/DWARF[4]), and ebisu dwarf (d2), which is defective in CYP90D2/D2 (ref. [5]). However, the grain yield of d61, brd1 and d2 mutants is decreased because of morphological alterations in their reproductive development.

===Function===

New cultivars with very erect leaves, which increase light capture for photosynthesis and nitrogen storage for grain filling, may have increased grain yields.The erect leaf phenotype of a rice brassinosteroid-deficient mutant, osdwarf4-1, is associated with enhanced grain yields under conditions of dense planting, even without extra fertilizer. Molecular and biochemical studies reveal that two different cytochrome P450s, CYP90B2/OsDWARF4 and CYP724B1/D11, function redundantly in C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis. Therefore, despite the central role of brassinosteroids in plant growth and development, mutation of OsDWARF4 alone causes only limited defects in brassinosteroid biosynthesis and plant morphology[6].

===Expression===
The affected gene, OsDWARF4, was identified by characterizing the site of insertion of the Tos17 retrotransposon.

----Molecular characterization of OsDWARF4 and OsDWARF4L1/D11

OsDWARF4 is a homolog of A. thaliana DWARF4, which encodes a cytochrome P450, CYP90B1 (Fig. 1a). CYP90B1/DWARF4 catalyzes C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis[7]. Only one DWARF4 homolog was identified in the rice genome. OsDWARF4 is located on the short arm of chromosome 3 (35 cM). Its predicted open reading frame consists of eight exons, and encodes a protein of 502 amino acids (Fig. 1b,c). OsDWARF4 is most closely related to DWARF4 (66.3% amino-acid identity; Fig. 1a). The structure of OsDWARF4 is similar to that of DWARF4 throughout its length: five domains found in cytochrome P450s—proline rich, A (also referred to as dioxygen binding), B (steroid binding), C and heme binding—are highly conserved (Fig. 1c). In wild-type rice, OsDWARF4 transcripts are most abundant in leaf blades and roots, but are also found in all other plant parts tested (Fig. 1d). Transcript levels were decreased by exogenous brassinolide treatment, and increased in brassinosteroid-insensitive d61-3 and brassinosteroid-deficient brd1-1 (Fig. 1e). These results suggest that OsDWARF4 expression is feedback regulated, as is that of A. thaliana brassinosteroid-biosynthesis genes[8].
OsDWARF4L1 was identified as an OsDWARF4-like gene by in silico screening of rice DNA databases. OsDWARF4L1 is located on the long arm of chromosome 4 (70.9 cM). A recent study demonstrated that a rice semi-dwarf and short-grain mutant, d11, has a loss-of-function mutation in the OsDWARF4L1 gene[9]. OsDWARF4L1/D11 shows the highest homology with A. thaliana CYP724A1, and is closely related to rice OsDWARF4 and A. thaliana DWARF4 (47.4%, 37.7% and 41.7% amino-acid identities, respectively; Fig. 1a). In wild-type rice, OsDWARF4L1/D11 transcripts were most abundant in vegetative shoot apices, leaf sheaths and roots, and found at more moderate levels in other organs (Fig. 1d). They were decreased by exogenous brassinolide treatment, and increased in d61-3 and brd1-1 (Fig. 1e). These results indicate that OsDWARF4L1/D11 expression is regulated by a feedback mechanism, as is that of OsDWARF4.

----Phenotypes of rice mutants defective in brassinosteroid biosynthesis

In osdwarf4-1, Tos17 insertion into exon 6 causes a premature termination in domain C (arrowhead in Fig. 1b,c). Because a premature stop codon before the heme-binding domain caused loss-of-function of A. thaliana DWARF4 (ref. [7]), we consider osdwarf4-1 to be a null allele. Unexpectedly, osdwarf4-1 plants show only weak morphological phenotypes, such as a slightly dwarfed stature and more erect leaves (Fig. 2a,b). Both of these phenotypes were complemented by the introduction of an entire OsDWARF4 gene (data not shown). These results seem inconsistent with the fact that rice d61-3 and brd1-1, which are loss-of-function mutants of OsBRI1 and OsDWARF respectively, display severe dwarfing and have malformed leaves (for example, see brd1-1 in Fig. 2a), and suggest functional redundancy in brassinosteroid biosynthesis in rice.
Because null alleles of d11, such as d11-4, also show a weak phenotype[9] (Fig. 2a),OsDWARF4L1/D11 and OsDWARF4 may function redundantly. To confirm this possibility, researcher produced double mutants by crossing homozygous osdwarf4-1 and d11-4 plants. Among the F2 population, 34 of 576 plants showed severe dwarfing and malformed leaves with tortuous leaf blades, indistinguishable from the brassinosteroid-deficient brd1-1 phenotype (Fig. 2a,c,d). This ratio fits the theoretical 9:3:3:1 (wild-type:osdwarf4-1:d11-4:osdwarf4-1/d11-4) segregation predicted by mendelian inheritance. PCR and sequence analyses confirmed that these plants were osdwarf4-1/d11-4 double mutants (data not shown).
Endogenous levels of brassinosteroid intermediates in brd1 clearly indicate that OsDWARF catalyzes C-6 oxidation[4]. However, quantification in d2 could not determine the step(s) catalyzed by CYP90D2/D2, because CYP90D2/D2 functions redundantly with another gene, CYP90D3, and loss-of-function of CYP90D2/D2 have caused only limited defects in brassinosteroid biosynthesis[5]. Similarly, endogenous levels of brassinosteroid intermediates were obviously changed in the osdwarf4-1/d11-4 double mutant relative to osdwarf4-1 or d11 single mutants. Although brassinolide was not detected in either the wild-type or osdwarf4-1/d11-4, another bioactive brassinosteroid, castasterone (CS), was detected in the wild-type but not in osdwarf4-1/d11-4, confirming that osdwarf4-1/d11-4 is brassinosteroid-deficient (Fig. 2e). Levels of the other five intermediates downstream of C-22 hydroxylation—6-deoxocathasterone (6-DeoxoCT), 6-deoxoteasterone (6-DeoxoTE), 3-dehydro-6-deoxoteasterone (6-Deoxo3DT), 6-deoxotyphasterol (6-DeoxoTY) and 6-deoxocastasterone (6-DeoxoCS)—were also greatly reduced in osdwarf4-1/d11-4 (Fig. 2e). This indicates that both proteins are involved in C-22 hydroxylation. It is unclear why the upstream compound, campestanol, did not accumulate in osdwarf4-1/d11-4. Because A. thaliana CYP90B1/DWARF4 also catalyzes C-22 hydroxylation of other upstream compounds such as campesterol and (24R)-ergost-4-en-3-one[10], it is possible that campestanol is not the major substrate for CYP90B2/OsDWARF4 and CYP724B1/D11 in rice.

----Characterization of a field-grown osdwarf4-1 mutant

Because the harvest index (grain/(grain plus straw)) of rice is about 0.5, further increases in yield potential will have to involve increases in crop biomass driven by more net photosynthesis[11]. In rice, the contribution of lower leaves to photosynthesis is significant, even though their photosynthetic capacity is considerably less than that of upper leaves[12]. Erect leaves allow greater penetration of light to lower leaves, thereby optimizing canopy photosynthesis[13]. Therefore, the most likely explanation for the increase in above-ground biomass in osdwarf4-1 plots under dense planting conditions (more than 1.2 times that in wild-type under conventional planting condition) is that the shade of the upper leaves is minimized, and the lower leaves receive more light to drive higher rates of photosynthesis (Fig. 3 and Supplementary Table 2 online). No difference was observed in the heading date (flowering time) and the grain-filling period (time to maturity) between wild-type and osdwarf4-1 plants (data not shown). In the wild-type, the above-ground biomass was increased in response to higher fertilizer application under conventional planting (22.2 plants m-2), but was significantly decreased under dense planting with doubled fertilizer application owing to reduced photosynthesis caused by lodging (Fig. 3a). In contrast, the above-ground biomass of osdwarf4-1 was significantly (P < 0.01) increased by dense planting at each fertilizer rate. Consequently, the above-ground biomass in osdwarf4-1 plots under dense planting with standard and a 1.5-fold increase in fertilizer application was approx1.34 and 1.52 times that in wild-type plots under conventional cultivation conditions. The increased biomass depended mainly on accelerated formation of tillers, which have the ability to generate a panicle (panicle number, Fig. 3b). Both total and fertile grain numbers tended to increase in densely planted osdwarf4-1 plots (Fig. 3c), resulting in significantly increased grain yield even when the fertilizer application was not increased (for example, the osdwarf4-1 yield was 6.19 t ha-1 under dense planting with standard fertilizer application whereas the wild-type yield was 4.69 t ha-1 under conventional conditions; Fig. 3d). However, the increase in grain yield was not as much as that in above-ground biomass, because the harvest index of osdwarf4-1 (0.41) was slightly lower than that of the wild-type (0.44). A similar field test gave comparable results.

----Overexpression of OsDWARF4

Overexpression of DWARF in transgenic tomato increases plant height[14], and overexpression of DWARF4 in transgenic A. thaliana and tobacco increases vegetative growth and seed yield[15]. In rice, however, the role of brassinosteroid in regulating leaf angle has been known for over 40 years[16, 17]. Whereas brassinosteroid deficiency increased the erectness of leaves, the opposite phenotype (increased leaf angle) was observed in transgenic rice overexpressing OsDWARF4 (27.6° versus 10.0° in the wild-type, P < 0.01), and was inherited in their progeny (data not shown). Because inclined leaves are unfavorable for photosynthesis when plants are grown in a group, we have to carefully evaluate whether the overexpression of a brassinosteroid-biosynthesis gene in transgenic rice increases grain yield as is the case in transgenic plants overexpressing DWARF4.

===Conclusion===

More erect leaves improve not only light distribution but also the size and use of leaf nitrogen reservoirs for grain growth[13]. The erect leaf phenotype caused by brassinosteroid deficiency improved biomass production under field conditions, resulting in increased grain yield without the negative environmental effects associated with fertilizers. Recently identified rice homologs of A. thaliana BAS1 that encodes the brassinosteroid catabolic enzyme, CYP734A1 (ref. [18]). Overexpression of these rice homologs in transgenic rice successfully induced the brassinosteroid-deficient phenotypes (unpublished results). Although ideally, expression of these genes should be restricted to particular tissues and organs such as leaf laminae, it should be possible to generate erect leaf varieties by the introduction of a single transgene. In conclusion, engineering more erect leaves modulating brassinosteroid metabolism can be combined with dense planting to improve rice grain yields.

==Labs working on this gene==
Please input related labs here.

==References==
<references>
<ref name="ref1">Sakamoto, T. & Matsuoka, M. Generating high-yielding varieties by genetic manipulation of plant architecture. Curr. Opin. Biotech. 15, 144–147 (2004).</ref>
<ref name="ref2">Li, J. & Chory, J. A putative leucine-rich repeat receptor kinase involved in brassinosteroid signal transduction. Cell 90, 929–938 (1997).</ref>
<ref name="ref3">Yamamuro, C. et al. Loss of function of a rice brassinosteroid insensitive1 homolog prevents internode elongation and bending of the lamina joint. Plant Cell 12, 1591–1606 (2000).</ref>

<ref name="ref4">Hong, Z. et al. Loss-of-function of a rice brassinosteroid biosynthetic enzyme, C-6 oxidase, prevents the organized arrangement and polar elongation of cells in the leaves and stem. Plant J. 32, 495–508 (2002).</ref>
<ref name="ref5">Hong, Z. et al. A rice brassinosteroid-deficient mutant, ebisu dwarf (d2), is caused by a loss of function of a new member of cytochrome P450. Plant Cell 15, 2900–2910 (2003).</ref>
<ref name="ref6">Sakamoto T;Morinaka Y;Ohnishi T;Sunohara H;Fujioka S;Ueguchi Tanaka M;Mizutani M;Sakata K;Takatsuto S;Yoshida S;Tanaka H;Kitano H;Matsuoka M Erect leaves caused by brassinosteroid deficiency increase biomass production and grain yield in rice Nature Biotechnology, 2006, 24(1): 105-109.</ref>
<ref name="ref7">Choe, S. et al. The DWF4 gene of Arabidopsis encodes a cytochrome P450 that mediates multiple 22alpha-hydroxylation steps in brassinosteroid biosynthesis. Plant Cell 10, 231–243 (1998).</ref>
<ref name="ref8">Bancos, S. et al. Regulation of transcript levels of the Arabidopsis cytochrome P450 genes involved in brassinosteroid biosynthesis. Plant Physiol. 130, 504–513 (2002).</ref>
<ref name="ref9">Tanabe, S. et al. A novel cytochrome P450 is implicated in brassinosteroid biosynthesis via the characterization of a rice dwarf mutant, dwarf11, with reduced seed length. Plant Cell 17, 776–790 (2005).</ref>
<ref name="ref10">Fujioka, S., Takatsuto, S. & Yoshida, S. An early C-22 oxidation branch in brassinosteroid biosynthetic pathway. Plant Physiol. 130, 930–939 (2002).</ref>
<ref name="ref11">Mann, C.C. Crop scientists seek a new revolution. Science 283, 310–314 (1999).</ref>
<ref name="ref12">Horton, P. Prospects for crop improvement through the genetic manipulation of photosynthesis: morphological and biochemical aspects of light capture. J. Exp. Bot. 51, 475–485 (2000).</ref>
<ref name="ref13">Sinclair, T.R. & Sheehy, J.E. Erect leaves and photosynthesis in rice. Science 283, 1455 (1999).</ref>
<ref name="ref14">Bishop, G.J. et al. The tomato DWARF enzyme catalyses C-6 oxidation in brassinosteroid biosynthesis. Proc. Natl. Acad. Sci. USA 96, 1761–1766 (1999).</ref>
<ref name="ref15">Choe, S. et al. Overexpression of DWARF4 in the brassinosteroid biosynthetic pathway results in increased vegetative growth and seed yield in Arabidopsis. Plant J. 26, 573–582 (2001).</ref>
<ref name="ref16">Maeda, E. Rate of lamina inclination in excised rice leaves. Physiol. Plant. 18, 813–827 (1965).</ref>
<ref name="ref17">Wada, K. et al. Brassinolide and homobrassinolide promotion of lamina inclination of rice seedlings. Plant Cell Physiol. 22, 323–325 (1981).</ref>
<ref name="ref18">Neff, M.M. et al. BAS1: A gene regulating brassinosteroid levels and light responsiveness in Arabidopsis. Proc. Natl. Acad. Sci. USA 96, 15316–15323 (1999).</ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os03g0227700|
Description = Cytochrome P450 family protein|
Version = NM_001055982.2 GI:297600576 GeneID:4332134|
Length = 6529 bp|
Definition = Oryza sativa Japonica Group Os03g0227700, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 3|Chromosome 3]]|
AP = Chromosome 3:6765109..6771637|
CDS = 6765109..6765374,6765543..6765867,6765971..6766123,6768937..6769188,6769337..6769429 ,6770222..6770300,6770398..6770504,6771392..6771637|
GCID = <gbrowseImage1>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</cdnaseq>|
AA = <aaseq>MAAMMASITSELLFFLPFILLALLTFYTTTVAKCHGGHWWRGGT TPAKRKRMNLPPGAAGWPLVGETFGYLRAHPATSVGRFMEQHIARYGKIYRSSLFGER TVVSADAGLNRYILQNEGRLFECSYPRSIGGILGKWSMLVLVGDPHREMRAISLNFLS SVRLRAVLLPEVERHTLLVLRAWPPSSTFSAQHQAKKFTFNLMAKNIMSMDPGEEETE RLRREYITFMKGVVSAPLNLPGTPYWKALKSRAAILGVIERKMEERVEKLSKEDASVE QDDLLGWALKQSNLSKEQILDLLLSLLFAGHETSSMALALAIFFLEGCPKAVQELREE HLGIARRQRLRGECKLSWEDYKEMVFTQCVINETLRLGNVVRFLHRKVIKDVHYKGYD IPSGWKILPVLAAVHLDSSLYEDPQRFNPWRWKSSGSSGGLAQSSSFMPYGGGTRLCA GSELAKLEMAVFLHHLVLNFRWELAEPDQAFVFPFVDFPKGLPIRVHRIAQDDEQE</aaseq>|
DNA = <dnaseqindica>1..266#435..759#863..1015#3829..4080#4229..4321#5114..5192#5290..5396#6284..6529#atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtcggtggtttcattactcggtacacgcacacagctatataagcacgcgcactgctcgcctcgccgtgaagcaagcttagctagtagctaggatatcgatcgatcggtaataataggctgacggcgagctggggggtggcgtgcggttgcgtgcgtgcgtgcatgcaggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaaggtacaacacacatcgtcctccattaagttgagcatgcatatatcaatcagctcgctctagcgatcaatatcaatgaaacgcgcgtttgcgtgcgtgcgtgcagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaaggtacgtactcgtagctactagctactccctccgtttcgaaatgtttgacgccgttgactttttatcacatatttgaccgttcgttttatttaaaaaaattaagtaattattaattattttcctatcatttgattcattgttaaatatatttttatgtaggcatataattttacatatctcacaaaagtttttaaataagacgaacggtcaaacatttgctaaaaagtcaacggtgtcaaatatttcgaaacggaggaagtagctagctaccaccccacccgtgcggattgatcgatcgatcgatgtccctctctgcctacacacaccctgcgccccggccgtacgcgcgcaaagtcgtgcgcgcatgagttttactccttccgtttcaaaaaaaaaacgacctagacaagtaatatgatgtaacctagaacaacgaatctatacaggagtatatccaaattcattgtcctagattacatcatatctttgtctatgttgccttttttaggaggagagagtactttacatatgagttatgaccggccggccaggttcctttcttcccctattccgtatgcatactataagcagctgcatataggcatatactactatacccagcttagctaaagcaaacaacttgcaacgacgatcatgtgcaggtccctgttgacagaggttagagatgtaaaccatttttgttgttttcattatctaaagagaaaacaagttactacgacgatcgagatgtacgcaaggggatcatatttatattggagaggagagagagatgatttgcgacttgtgagtgataagtgaatatatatactatatatttgtgtaggagtgacgatgacttccctgtcaatgttttgtttggtatgcacacggggggaaaagagtttagagaaaggaattggctagagagggccaggtttagtttctttaatttttgtttggtggttttgtgtgtagctagtccgcatgcatgatatatcatatatatgcatggcccatcattagatatatatacatgacggcagtgcacttattatatatatgtaggggcatatacaaggtgtatgtgtttagaattgtggatggtccaggaattgttgcagctagttctatactagcatagtgaattagtgatccatatcgagatcaggtccaatcggtcagctcgcacagcgggtcttttagtacgatttgttcggacaaattatcttgcctttgagataatcaacataggattgaaactgcatactagcaatagaccatagctatttggcaatatagaaagctaaaaaatgttgattgaaaggtgcgtgcatgcatgctagcagtatgtcttttaggccaggtccagtttttttttttttctcaaaggcagttgcttaaggtgaaattctagcagaggtcatatgaatatatttattatcttccaaacatgagccaaaacctatatatagttaactcaaaaaaccatgcattggtacttcttctgagtatgcatcctggggactgatcgatcatgcagattcttttctttctttttccctggggacgtactgatcatcactcatgtgagcataaaatgttgcctactttttactgcatgtgaggtcacactcaaatgatctttattgattgcctcagcatgcatgctgctagctttcacgatctctctccgatacacacaccctgtgctgtgctcaacagcatgcaacctggcaggtggaatttttaatctgttggaaagtaaatgaccaaccttcataatttggcatgattatatgctggggaatctatcatttatcatgattcaaccgctgaccaaactctttgccacacacccgaaccacattaatttcgttaagcggatgattagccagcaaccttcacagtttggttttaattagttaatgaaagggtgcttgtagttttattagctgaagagacactgcagtactcctactgagcgttctccttaattgggaaatagatctctgcagtactatcctttaactttaagtgaataacatgtagctaggtctcacacttatcaggcccacacatcagtgatccaagtcaaatggcaatacaccataggagattcttccactaattggctaacaaaaataaactgtgatcactgatcacagctgctaacactataacacattaatttgcatgtctcatgaaatgccagatgagttggattaattatctatcttatgttcagtgccgaggaatatacaaggaaacatcattggtatagttgtggagttctgagtgcattatctcacccagtacggtcaaattaatcctagtctccgtaaatatggttgtgtgcactcttagttaattagtgtagacccggaaattatactttgtcgagaatgttgttatccgtgtgttaagtttcaagtaccctgcaggcctataactaaacactgccaacatggatgtttaagaatgaattatacatacagcgcacttggaaaagaaaaggggtagctacccagctgaagaacacctctctctcatttgaaggtcgacacaatatgtgtcttttctatatcgaaaggaaacgacattttttctcttcaaaacagaaagctatgacaagagacataatcctagactagtaggcggttaggcaacaccatatctgaatttgataggtgtcccacatattttttttcttttcctgataagaaggaacttttcaccgtgagcttatttttttctgttctatgtcgacattagttccatgatgtttactgtgtatttttactgttcccctgcagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggtacagagaattacagattcgaaattaacatctactgtcactagctcagtgctacgtttaactgtagagcaacaattgcagaaagcgaaggcatcagaaatgacatgggaatcgattagttacactcacactgacgcgtgacatgcaggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtaaggccccccctactttgttccttccgatccctcctccctaaacgctactaatataaatcggtgcaaaggctttgtgctagcgtagctcttgctgttttgtttggtcacttgcgtagagagatagcaacttcccgaaatgtcatgtacgaatggagactgaaagctctgcgaaataattgccgggccacgtactacagtacttacgctccctctcttcagtttggatcgatcgagctaagaaggcgcacgcatgtgtgcattgatcgatagagtcgatctgttctgccttgtaagctagccagctgaggttctttcgccttggaagtttctacagggtgaggcccatcgtcttgcaacgatccaaatgccatgacaagtacctgttgttgtaggcttccagtagcacggcctgcccccaagaaggtatctgaatcttctgcattgctgttcagacggcagagcagaacatgccgaaataccattccagtcagttgcagtatatgtgtgtggtggtatcagataaaggagcagagtaatttgaggccttagaaattcagatgagttgctcgacaattgtcgcc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Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001055982.2 RefSeq:Os03g0227700]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 3]]
[[Category:Chromosome 3]]

File:Figure 1 - Molecular characterization of OsDWARF4 and OsDWARF4L1-D11..jpg

2014-05-26T12:38:40Z

LQ: (a) Phylogenetic relationships among brassinosteroid-biosynthetic cytochrome P450s of rice, A. thaliana and tomato. Bar, 0.1 amino-acid substitutions per site. (b) Genomic organization of the OsDWARF4 gene. Boxes represent exons, and lines separating

File:Figure 1 - Molecular characterization of OsDWARF4 and OsDWARF4L1-D11..jpg

2014-05-26T12:37:53Z

LQ: uploaded a new version of "File:Figure 1 - Molecular characterization of OsDWARF4 and OsDWARF4L1-D11..jpg": (a) Phylogenetic relationships among brassinosteroid-biosynthetic cytochrome P450s of rice, A. thaliana and tomato. Bar, 0.1 amino-ac

(a) Phylogenetic relationships among brassinosteroid-biosynthetic cytochrome P450s of rice, A. thaliana and tomato. Bar, 0.1 amino-acid substitutions per site. (b) Genomic organization of the OsDWARF4 gene. Boxes represent exons, and lines separating boxes represent introns. The arrowhead indicates the site of Tos17 insertion in osdwarf4-1. (c) Relative positions of the major domains in OsDWARF4. All of the major domains found in the cytochrome P450 superfamily are conserved in OsDWARF4. The arrowhead indicates the site of Tos17 insertion in osdwarf4-1. (d) Expression of OsDWARF4 and OsDWARF4L1/D11 in various organs of wild-type rice. The gene encoding Histone H3 was used as a control. (e) Feedback regulation of OsDWARF4 and OsDWARF4L1/D11 in brassinolide-treated wild-type plants (left) and brassinosteroid-related mutants (right). Values under each panel indicate relative amounts of each transcript. (f) Function of recombinant CYP90B2/OsDWARF4 and CYP724B1/D11 proteins. Campesterol (CR) was incubated with each recombinant protein and analyzed by GC-MS. Inset shows that the mass spectrum of the trimethylsilyl ester of each metabolite coincided with that of the trimethylsilyl ether of synthesized (22S)-22-hydroxycampesterol (22-OHCR).

File:Figure 1 - Molecular characterization of OsDWARF4 and OsDWARF4L1-D11..jpg

2014-05-26T08:54:18Z

File talk:Figure 1 - Molecular characterization of OsDWARF4 and OsDWARF4L1-D11..jpg

2014-05-26T08:40:30Z

LQ: Created page with "(a) Phylogenetic relationships among brassinosteroid-biosynthetic cytochrome P450s of rice, A. thaliana and tomato. Bar, 0.1 amino-acid substitutions per site. (b) Genomic org..."

File:Figure 1 - Molecular characterization of OsDWARF4 and OsDWARF4L1-D11..jpg

2014-05-26T08:39:39Z

File:Figure 1 - Molecular characterization of OsDWARF4 and OsDWARF4L1-D11..jpg

2014-05-26T08:33:24Z

Os03g0227700

2014-05-24T13:37:25Z

LQ: /* References */

Please input one-sentence summary here.
''OsDWAR''F4 is expressed in young elongating tissues.
==Annotated Information==

Brassinosteroids influence both plant height and leaf erectness in rice[1]. Arabidopsis thaliana BRASSINOSTEROID INSENSITIVE1 (bri1) mutants, which are deficient in the brassinosteroid receptor, have a severe dwarf phenotype[2]. The rice mutants d61-1 and d61-2, which are weak mutant alleles of OsBRI1, a homolog of A. thaliana BRI1, have both a semi-dwarf phenotype and more erect leaves[3]. In rice, an erect leaf phenotype has also been associated with loss-of-function, brassinosteroid-deficient mutants such as brassinosteroid-deficient dwarf1 (brd1), which is defective in OsDWARF (the homolog of tomato CYP85A1/DWARF[4]), and ebisu dwarf (d2), which is defective in CYP90D2/D2 (ref. [5]). However, the grain yield of d61, brd1 and d2 mutants is decreased because of morphological alterations in their reproductive development.

===Function===

New cultivars with very erect leaves, which increase light capture for photosynthesis and nitrogen storage for grain filling, may have increased grain yields.The erect leaf phenotype of a rice brassinosteroid-deficient mutant, osdwarf4-1, is associated with enhanced grain yields under conditions of dense planting, even without extra fertilizer. Molecular and biochemical studies reveal that two different cytochrome P450s, CYP90B2/OsDWARF4 and CYP724B1/D11, function redundantly in C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis. Therefore, despite the central role of brassinosteroids in plant growth and development, mutation of OsDWARF4 alone causes only limited defects in brassinosteroid biosynthesis and plant morphology[6].

===Expression===
The affected gene, OsDWARF4, was identified by characterizing the site of insertion of the Tos17 retrotransposon.

----Molecular characterization of OsDWARF4 and OsDWARF4L1/D11

OsDWARF4 is a homolog of A. thaliana DWARF4, which encodes a cytochrome P450, CYP90B1 (Fig. 1a). CYP90B1/DWARF4 catalyzes C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis[7]. Only one DWARF4 homolog was identified in the rice genome. OsDWARF4 is located on the short arm of chromosome 3 (35 cM). Its predicted open reading frame consists of eight exons, and encodes a protein of 502 amino acids (Fig. 1b,c). OsDWARF4 is most closely related to DWARF4 (66.3% amino-acid identity; Fig. 1a). The structure of OsDWARF4 is similar to that of DWARF4 throughout its length: five domains found in cytochrome P450s—proline rich, A (also referred to as dioxygen binding), B (steroid binding), C and heme binding—are highly conserved (Fig. 1c). In wild-type rice, OsDWARF4 transcripts are most abundant in leaf blades and roots, but are also found in all other plant parts tested (Fig. 1d). Transcript levels were decreased by exogenous brassinolide treatment, and increased in brassinosteroid-insensitive d61-3 and brassinosteroid-deficient brd1-1 (Fig. 1e). These results suggest that OsDWARF4 expression is feedback regulated, as is that of A. thaliana brassinosteroid-biosynthesis genes[8].
OsDWARF4L1 was identified as an OsDWARF4-like gene by in silico screening of rice DNA databases. OsDWARF4L1 is located on the long arm of chromosome 4 (70.9 cM). A recent study demonstrated that a rice semi-dwarf and short-grain mutant, d11, has a loss-of-function mutation in the OsDWARF4L1 gene[9]. OsDWARF4L1/D11 shows the highest homology with A. thaliana CYP724A1, and is closely related to rice OsDWARF4 and A. thaliana DWARF4 (47.4%, 37.7% and 41.7% amino-acid identities, respectively; Fig. 1a). In wild-type rice, OsDWARF4L1/D11 transcripts were most abundant in vegetative shoot apices, leaf sheaths and roots, and found at more moderate levels in other organs (Fig. 1d). They were decreased by exogenous brassinolide treatment, and increased in d61-3 and brd1-1 (Fig. 1e). These results indicate that OsDWARF4L1/D11 expression is regulated by a feedback mechanism, as is that of OsDWARF4.

----Phenotypes of rice mutants defective in brassinosteroid biosynthesis

In osdwarf4-1, Tos17 insertion into exon 6 causes a premature termination in domain C (arrowhead in Fig. 1b,c). Because a premature stop codon before the heme-binding domain caused loss-of-function of A. thaliana DWARF4 (ref. [7]), we consider osdwarf4-1 to be a null allele. Unexpectedly, osdwarf4-1 plants show only weak morphological phenotypes, such as a slightly dwarfed stature and more erect leaves (Fig. 2a,b). Both of these phenotypes were complemented by the introduction of an entire OsDWARF4 gene (data not shown). These results seem inconsistent with the fact that rice d61-3 and brd1-1, which are loss-of-function mutants of OsBRI1 and OsDWARF respectively, display severe dwarfing and have malformed leaves (for example, see brd1-1 in Fig. 2a), and suggest functional redundancy in brassinosteroid biosynthesis in rice.
Because null alleles of d11, such as d11-4, also show a weak phenotype[9] (Fig. 2a),OsDWARF4L1/D11 and OsDWARF4 may function redundantly. To confirm this possibility, researcher produced double mutants by crossing homozygous osdwarf4-1 and d11-4 plants. Among the F2 population, 34 of 576 plants showed severe dwarfing and malformed leaves with tortuous leaf blades, indistinguishable from the brassinosteroid-deficient brd1-1 phenotype (Fig. 2a,c,d). This ratio fits the theoretical 9:3:3:1 (wild-type:osdwarf4-1:d11-4:osdwarf4-1/d11-4) segregation predicted by mendelian inheritance. PCR and sequence analyses confirmed that these plants were osdwarf4-1/d11-4 double mutants (data not shown).
Endogenous levels of brassinosteroid intermediates in brd1 clearly indicate that OsDWARF catalyzes C-6 oxidation[4]. However, quantification in d2 could not determine the step(s) catalyzed by CYP90D2/D2, because CYP90D2/D2 functions redundantly with another gene, CYP90D3, and loss-of-function of CYP90D2/D2 have caused only limited defects in brassinosteroid biosynthesis[5]. Similarly, endogenous levels of brassinosteroid intermediates were obviously changed in the osdwarf4-1/d11-4 double mutant relative to osdwarf4-1 or d11 single mutants. Although brassinolide was not detected in either the wild-type or osdwarf4-1/d11-4, another bioactive brassinosteroid, castasterone (CS), was detected in the wild-type but not in osdwarf4-1/d11-4, confirming that osdwarf4-1/d11-4 is brassinosteroid-deficient (Fig. 2e). Levels of the other five intermediates downstream of C-22 hydroxylation—6-deoxocathasterone (6-DeoxoCT), 6-deoxoteasterone (6-DeoxoTE), 3-dehydro-6-deoxoteasterone (6-Deoxo3DT), 6-deoxotyphasterol (6-DeoxoTY) and 6-deoxocastasterone (6-DeoxoCS)—were also greatly reduced in osdwarf4-1/d11-4 (Fig. 2e). This indicates that both proteins are involved in C-22 hydroxylation. It is unclear why the upstream compound, campestanol, did not accumulate in osdwarf4-1/d11-4. Because A. thaliana CYP90B1/DWARF4 also catalyzes C-22 hydroxylation of other upstream compounds such as campesterol and (24R)-ergost-4-en-3-one[10], it is possible that campestanol is not the major substrate for CYP90B2/OsDWARF4 and CYP724B1/D11 in rice.

----Characterization of a field-grown osdwarf4-1 mutant

Because the harvest index (grain/(grain plus straw)) of rice is about 0.5, further increases in yield potential will have to involve increases in crop biomass driven by more net photosynthesis[11]. In rice, the contribution of lower leaves to photosynthesis is significant, even though their photosynthetic capacity is considerably less than that of upper leaves[12]. Erect leaves allow greater penetration of light to lower leaves, thereby optimizing canopy photosynthesis[13]. Therefore, the most likely explanation for the increase in above-ground biomass in osdwarf4-1 plots under dense planting conditions (more than 1.2 times that in wild-type under conventional planting condition) is that the shade of the upper leaves is minimized, and the lower leaves receive more light to drive higher rates of photosynthesis (Fig. 3 and Supplementary Table 2 online). No difference was observed in the heading date (flowering time) and the grain-filling period (time to maturity) between wild-type and osdwarf4-1 plants (data not shown). In the wild-type, the above-ground biomass was increased in response to higher fertilizer application under conventional planting (22.2 plants m-2), but was significantly decreased under dense planting with doubled fertilizer application owing to reduced photosynthesis caused by lodging (Fig. 3a). In contrast, the above-ground biomass of osdwarf4-1 was significantly (P < 0.01) increased by dense planting at each fertilizer rate. Consequently, the above-ground biomass in osdwarf4-1 plots under dense planting with standard and a 1.5-fold increase in fertilizer application was approx1.34 and 1.52 times that in wild-type plots under conventional cultivation conditions. The increased biomass depended mainly on accelerated formation of tillers, which have the ability to generate a panicle (panicle number, Fig. 3b). Both total and fertile grain numbers tended to increase in densely planted osdwarf4-1 plots (Fig. 3c), resulting in significantly increased grain yield even when the fertilizer application was not increased (for example, the osdwarf4-1 yield was 6.19 t ha-1 under dense planting with standard fertilizer application whereas the wild-type yield was 4.69 t ha-1 under conventional conditions; Fig. 3d). However, the increase in grain yield was not as much as that in above-ground biomass, because the harvest index of osdwarf4-1 (0.41) was slightly lower than that of the wild-type (0.44). A similar field test gave comparable results.

----Overexpression of OsDWARF4

Overexpression of DWARF in transgenic tomato increases plant height[14], and overexpression of DWARF4 in transgenic A. thaliana and tobacco increases vegetative growth and seed yield[15]. In rice, however, the role of brassinosteroid in regulating leaf angle has been known for over 40 years[16, 17]. Whereas brassinosteroid deficiency increased the erectness of leaves, the opposite phenotype (increased leaf angle) was observed in transgenic rice overexpressing OsDWARF4 (27.6° versus 10.0° in the wild-type, P < 0.01), and was inherited in their progeny (data not shown). Because inclined leaves are unfavorable for photosynthesis when plants are grown in a group, we have to carefully evaluate whether the overexpression of a brassinosteroid-biosynthesis gene in transgenic rice increases grain yield as is the case in transgenic plants overexpressing DWARF4.

===Conclusion===

More erect leaves improve not only light distribution but also the size and use of leaf nitrogen reservoirs for grain growth[13]. The erect leaf phenotype caused by brassinosteroid deficiency improved biomass production under field conditions, resulting in increased grain yield without the negative environmental effects associated with fertilizers. Recently identified rice homologs of A. thaliana BAS1 that encodes the brassinosteroid catabolic enzyme, CYP734A1 (ref. [18]). Overexpression of these rice homologs in transgenic rice successfully induced the brassinosteroid-deficient phenotypes (unpublished results). Although ideally, expression of these genes should be restricted to particular tissues and organs such as leaf laminae, it should be possible to generate erect leaf varieties by the introduction of a single transgene. In conclusion, engineering more erect leaves modulating brassinosteroid metabolism can be combined with dense planting to improve rice grain yields.

==Labs working on this gene==
Please input related labs here.

==References==
[1] Sakamoto, T. & Matsuoka, M. Generating high-yielding varieties by genetic manipulation of plant architecture. Curr. Opin. Biotech. 15, 144–147 (2004).
[2] Li, J. & Chory, J. A putative leucine-rich repeat receptor kinase involved in brassinosteroid signal transduction. Cell 90, 929–938 (1997).
[3] Yamamuro, C. et al. Loss of function of a rice brassinosteroid insensitive1 homolog prevents internode elongation and bending of the lamina joint. Plant Cell 12, 1591–1606 (2000).
[4] Hong, Z. et al. Loss-of-function of a rice brassinosteroid biosynthetic enzyme, C-6 oxidase, prevents the organized arrangement and polar elongation of cells in the leaves and stem. Plant J. 32, 495–508 (2002).
[5] Hong, Z. et al. A rice brassinosteroid-deficient mutant, ebisu dwarf (d2), is caused by a loss of function of a new member of cytochrome P450. Plant Cell 15, 2900–2910 (2003).
[6]Sakamoto T;Morinaka Y;Ohnishi T;Sunohara H;Fujioka S;Ueguchi Tanaka M;Mizutani M;Sakata K;Takatsuto S;Yoshida S;Tanaka H;Kitano H;Matsuoka M Erect leaves caused by brassinosteroid deficiency increase biomass production and grain yield in rice Nature Biotechnology, 2006, 24(1): 105-109.
[7] Choe, S. et al. The DWF4 gene of Arabidopsis encodes a cytochrome P450 that mediates multiple 22alpha-hydroxylation steps in brassinosteroid biosynthesis. Plant Cell 10, 231–243 (1998).
[8] Bancos, S. et al. Regulation of transcript levels of the Arabidopsis cytochrome P450 genes involved in brassinosteroid biosynthesis. Plant Physiol. 130, 504–513 (2002).
[9] Tanabe, S. et al. A novel cytochrome P450 is implicated in brassinosteroid biosynthesis via the characterization of a rice dwarf mutant, dwarf11, with reduced seed length. Plant Cell 17, 776–790 (2005).
[10] Fujioka, S., Takatsuto, S. & Yoshida, S. An early C-22 oxidation branch in brassinosteroid biosynthetic pathway. Plant Physiol. 130, 930–939 (2002).
[11] Mann, C.C. Crop scientists seek a new revolution. Science 283, 310–314 (1999).
[12] Horton, P. Prospects for crop improvement through the genetic manipulation of photosynthesis: morphological and biochemical aspects of light capture. J. Exp. Bot. 51, 475–485 (2000).
[13] Sinclair, T.R. & Sheehy, J.E. Erect leaves and photosynthesis in rice. Science 283, 1455 (1999).
[14] Bishop, G.J. et al. The tomato DWARF enzyme catalyses C-6 oxidation in brassinosteroid biosynthesis. Proc. Natl. Acad. Sci. USA 96, 1761–1766 (1999).
[15] Choe, S. et al. Overexpression of DWARF4 in the brassinosteroid biosynthetic pathway results in increased vegetative growth and seed yield in Arabidopsis. Plant J. 26, 573–582 (2001).
[16] Maeda, E. Rate of lamina inclination in excised rice leaves. Physiol. Plant. 18, 813–827 (1965).
[17] Wada, K. et al. Brassinolide and homobrassinolide promotion of lamina inclination of rice seedlings. Plant Cell Physiol. 22, 323–325 (1981).
[18] Neff, M.M. et al. BAS1: A gene regulating brassinosteroid levels and light responsiveness in Arabidopsis. Proc. Natl. Acad. Sci. USA 96, 15316–15323 (1999).

==Structured Information==
{{JaponicaGene|
GeneName = Os03g0227700|
Description = Cytochrome P450 family protein|
Version = NM_001055982.2 GI:297600576 GeneID:4332134|
Length = 6529 bp|
Definition = Oryza sativa Japonica Group Os03g0227700, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 3|Chromosome 3]]|
AP = Chromosome 3:6765109..6771637|
CDS = 6765109..6765374,6765543..6765867,6765971..6766123,6768937..6769188,6769337..6769429 ,6770222..6770300,6770398..6770504,6771392..6771637|
GCID = <gbrowseImage1>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</cdnaseq>|
AA = <aaseq>MAAMMASITSELLFFLPFILLALLTFYTTTVAKCHGGHWWRGGT TPAKRKRMNLPPGAAGWPLVGETFGYLRAHPATSVGRFMEQHIARYGKIYRSSLFGER TVVSADAGLNRYILQNEGRLFECSYPRSIGGILGKWSMLVLVGDPHREMRAISLNFLS SVRLRAVLLPEVERHTLLVLRAWPPSSTFSAQHQAKKFTFNLMAKNIMSMDPGEEETE RLRREYITFMKGVVSAPLNLPGTPYWKALKSRAAILGVIERKMEERVEKLSKEDASVE QDDLLGWALKQSNLSKEQILDLLLSLLFAGHETSSMALALAIFFLEGCPKAVQELREE HLGIARRQRLRGECKLSWEDYKEMVFTQCVINETLRLGNVVRFLHRKVIKDVHYKGYD IPSGWKILPVLAAVHLDSSLYEDPQRFNPWRWKSSGSSGGLAQSSSFMPYGGGTRLCA GSELAKLEMAVFLHHLVLNFRWELAEPDQAFVFPFVDFPKGLPIRVHRIAQDDEQE</aaseq>|
DNA = <dnaseqindica>1..266#435..759#863..1015#3829..4080#4229..4321#5114..5192#5290..5396#6284..6529#atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtcggtggtttcattactcggtacacgcacacagctatataagcacgcgcactgctcgcctcgccgtgaagcaagcttagctagtagctaggatatcgatcgatcggtaataataggctgacggcgagctggggggtggcgtgcggttgcgtgcgtgcgtgcatgcaggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaaggtacaacacacatcgtcctccattaagttgagcatgcatatatcaatcagctcgctctagcgatcaatatcaatgaaacgcgcgtttgcgtgcgtgcgtgcagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaaggtacgtactcgtagctactagctactccctccgtttcgaaatgtttgacgccgttgactttttatcacatatttgaccgttcgttttatttaaaaaaattaagtaattattaattattttcctatcatttgattcattgttaaatatatttttatgtaggcatataattttacatatctcacaaaagtttttaaataagacgaacggtcaaacatttgctaaaaagtcaacggtgtcaaatatttcgaaacggaggaagtagctagctaccaccccacccgtgcggattgatcgatcgatcgatgtccctctctgcctacacacaccctgcgccccggccgtacgcgcgcaaagtcgtgcgcgcatgagttttactccttccgtttcaaaaaaaaaacgacctagacaagtaatatgatgtaacctagaacaacgaatctatacaggagtatatccaaattcattgtcctagattacatcatatctttgtctatgttgccttttttaggaggagagagtactttacatatgagttatgaccggccggccaggttcctttcttcccctattccgtatgcatactataagcagctgcatataggcatatactactatacccagcttagctaaagcaaacaacttgcaacgacgatcatgtgcaggtccctgttgacagaggttagagatgtaaaccatttttgttgttttcattatctaaagagaaaacaagttactacgacgatcgagatgtacgcaaggggatcatatttatattggagaggagagagagatgatttgcgacttgtgagtgataagtgaatatatatactatatatttgtgtaggagtgacgatgacttccctgtcaatgttttgtttggtatgcacacggggggaaaagagtttagagaaaggaattggctagagagggccaggtttagtttctttaatttttgtttggtggttttgtgtgtagctagtccgcatgcatgatatatcatatatatgcatggcccatcattagatatatatacatgacggcagtgcacttattatatatatgtaggggcatatacaaggtgtatgtgtttagaattgtggatggtccaggaattgttgcagctagttctatactagcatagtgaattagtgatccatatcgagatcaggtccaatcggtcagctcgcacagcgggtcttttagtacgatttgttcggacaaattatcttgcctttgagataatcaacataggattgaaactgcatactagcaatagaccatagctatttggcaatatagaaagctaaaaaatgttgattgaaaggtgcgtgcatgcatgctagcagtatgtcttttaggccaggtccagtttttttttttttctcaaaggcagttgcttaaggtgaaattctagcagaggtcatatgaatatatttattatcttccaaacatgagccaaaacctatatatagttaactcaaaaaaccatgcattggtacttcttctgagtatgcatcctggggactgatcgatcatgcagattcttttctttctttttccctggggacgtactgatcatcactcatgtgagcataaaatgttgcctactttttactgcatgtgaggtcacactcaaatgatctttattgattgcctcagcatgcatgctgctagctttcacgatctctctccgatacacacaccctgtgctgtgctcaacagcatgcaacctggcaggtggaatttttaatctgttggaaagtaaatgaccaaccttcataatttggcatgattatatgctggggaatctatcatttatcatgattcaaccgctgaccaaactctttgccacacacccgaaccacattaatttcgttaagcggatgattagccagcaaccttcacagtttggttttaattagttaatgaaagggtgcttgtagttttattagctgaagagacactgcagtactcctactgagcgttctccttaattgggaaatagatctctgcagtactatcctttaactttaagtgaataacatgtagctaggtctcacacttatcaggcccacacatcagtgatccaagtcaaatggcaatacaccataggagattcttccactaattggctaacaaaaataaactgtgatcactgatcacagctgctaacactataacacattaatttgcatgtctcatgaaatgccagatgagttggattaattatctatcttatgttcagtgccgaggaatatacaaggaaacatcattggtatagttgtggagttctgagtgcattatctcacccagtacggtcaaattaatcctagtctccgtaaatatggttgtgtgcactcttagttaattagtgtagacccggaaattatactttgtcgagaatgttgttatccgtgtgttaagtttcaagtaccctgcaggcctataactaaacactgccaacatggatgtttaagaatgaattatacatacagcgcacttggaaaagaaaaggggtagctacccagctgaagaacacctctctctcatttgaaggtcgacacaatatgtgtcttttctatatcgaaaggaaacgacattttttctcttcaaaacagaaagctatgacaagagacataatcctagactagtaggcggttaggcaacaccatatctgaatttgataggtgtcccacatattttttttcttttcctgataagaaggaacttttcaccgtgagcttatttttttctgttctatgtcgacattagttccatgatgtttactgtgtatttttactgttcccctgcagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggtacagagaattacagattcgaaattaacatctactgtcactagctcagtgctacgtttaactgtagagcaacaattgcagaaagcgaaggcatcagaaatgacatgggaatcgattagttacactcacactgacgcgtgacatgcaggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtaaggccccccctactttgttccttccgatccctcctccctaaacgctactaatataaatcggtgcaaaggctttgtgctagcgtagctcttgctgttttgtttggtcacttgcgtagagagatagcaacttcccgaaatgtcatgtacgaatggagactgaaagctctgcgaaataattgccgggccacgtactacagtacttacgctccctctcttcagtttggatcgatcgagctaagaaggcgcacgcatgtgtgcattgatcgatagagtcgatctgttctgccttgtaagctagccagctgaggttctttcgccttggaagtttctacagggtgaggcccatcgtcttgcaacgatccaaatgccatgacaagtacctgttgttgtaggcttccagtagcacggcctgcccccaagaaggtatctgaatcttctgcattgctgttcagacggcagagcagaacatgccgaaataccattccagtcagttgcagtatatgtgtgtggtggtatcagataaaggagcagagtaatttgaggccttagaaattcagatgagttgctcgacaattgtcgcctaccgatcgagaataagttgcatgcatgtatgtgatgtaattatgggtcttagtcagcaggatagtttagtcgctgcagtactgtagattgtagtacttgcgttcaaagcagacccaaaacagaacttgcattcaggaaacaaaacatgaaaaaagaaatatacagtacttggcactgatatttcttttgtttgggtccttattgcaggtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggtgagtagctcgatttcactcttcttccctgaatcgacctctcctctatctgacagcagaagctaactgaactgaactgaactgaactgaactgaaggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaaggtcagtcgcagtaggattatcagtgaatctaatctattaatctcggtttaattgctgggattcaaggaaaagaaaatactgggacaagcgcggatgtcgctgttgggcagtgtctgtcaatcataagcgaaggggatcatgtcagaatctgcaaaagcagcacaggctaaacgcgtgcttgcctcgagcagcaatcatgcagagaggcaagctccgcgataggaacatgcatggtgaatcactgaacctgatatgaaagtgaatgctcagctcagctcgctcgctcgcgcgcgcatgtgctttctcatttcgccgtgttgcgtgctagctagtcaatggtgttcccacgaattgttttctcttctcatcacacaaagtcagtggtgaaaaccgaggtgcatgtgcggtaatggaacgggggattaggcggcgacgccgtgatggattatgcattcgtgcgagaaaaatccaggagcgttggtccaaatcgcaagcggatcaaaccgaaccaatcagtggacgattggagccctgcagtccatccatcacatctccattcttatccgctgcgactcacggacattagttgttgaggggtgaacgacgcatgtgtgtatgtatgctcgatcgatcgatcgaacaatggatctgtgtgtggtgtagttgccaacgccgacgagattatctgctcgtctgcatgcagccgcgaaaaagacatgtgtccggattatcctgcaggagattgagcaagataatctgctgcgcaaaaactgcagccaatcgatcattccatgacgcgaaaaaaaaaacaataattactactgctgatacgataacttcgcttcgaattgttgacacacaagtgttcttcttttgttcgattgcagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001055982.2 RefSeq:Os03g0227700]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 3]]
[[Category:Chromosome 3]]

Os03g0227700

2014-05-24T13:36:41Z

LQ: /* References */

Please input one-sentence summary here.
''OsDWAR''F4 is expressed in young elongating tissues.
==Annotated Information==

Brassinosteroids influence both plant height and leaf erectness in rice[1]. Arabidopsis thaliana BRASSINOSTEROID INSENSITIVE1 (bri1) mutants, which are deficient in the brassinosteroid receptor, have a severe dwarf phenotype[2]. The rice mutants d61-1 and d61-2, which are weak mutant alleles of OsBRI1, a homolog of A. thaliana BRI1, have both a semi-dwarf phenotype and more erect leaves[3]. In rice, an erect leaf phenotype has also been associated with loss-of-function, brassinosteroid-deficient mutants such as brassinosteroid-deficient dwarf1 (brd1), which is defective in OsDWARF (the homolog of tomato CYP85A1/DWARF[4]), and ebisu dwarf (d2), which is defective in CYP90D2/D2 (ref. [5]). However, the grain yield of d61, brd1 and d2 mutants is decreased because of morphological alterations in their reproductive development.

===Function===

New cultivars with very erect leaves, which increase light capture for photosynthesis and nitrogen storage for grain filling, may have increased grain yields.The erect leaf phenotype of a rice brassinosteroid-deficient mutant, osdwarf4-1, is associated with enhanced grain yields under conditions of dense planting, even without extra fertilizer. Molecular and biochemical studies reveal that two different cytochrome P450s, CYP90B2/OsDWARF4 and CYP724B1/D11, function redundantly in C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis. Therefore, despite the central role of brassinosteroids in plant growth and development, mutation of OsDWARF4 alone causes only limited defects in brassinosteroid biosynthesis and plant morphology[6].

===Expression===
The affected gene, OsDWARF4, was identified by characterizing the site of insertion of the Tos17 retrotransposon.

----Molecular characterization of OsDWARF4 and OsDWARF4L1/D11

OsDWARF4 is a homolog of A. thaliana DWARF4, which encodes a cytochrome P450, CYP90B1 (Fig. 1a). CYP90B1/DWARF4 catalyzes C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis[7]. Only one DWARF4 homolog was identified in the rice genome. OsDWARF4 is located on the short arm of chromosome 3 (35 cM). Its predicted open reading frame consists of eight exons, and encodes a protein of 502 amino acids (Fig. 1b,c). OsDWARF4 is most closely related to DWARF4 (66.3% amino-acid identity; Fig. 1a). The structure of OsDWARF4 is similar to that of DWARF4 throughout its length: five domains found in cytochrome P450s—proline rich, A (also referred to as dioxygen binding), B (steroid binding), C and heme binding—are highly conserved (Fig. 1c). In wild-type rice, OsDWARF4 transcripts are most abundant in leaf blades and roots, but are also found in all other plant parts tested (Fig. 1d). Transcript levels were decreased by exogenous brassinolide treatment, and increased in brassinosteroid-insensitive d61-3 and brassinosteroid-deficient brd1-1 (Fig. 1e). These results suggest that OsDWARF4 expression is feedback regulated, as is that of A. thaliana brassinosteroid-biosynthesis genes[8].
OsDWARF4L1 was identified as an OsDWARF4-like gene by in silico screening of rice DNA databases. OsDWARF4L1 is located on the long arm of chromosome 4 (70.9 cM). A recent study demonstrated that a rice semi-dwarf and short-grain mutant, d11, has a loss-of-function mutation in the OsDWARF4L1 gene[9]. OsDWARF4L1/D11 shows the highest homology with A. thaliana CYP724A1, and is closely related to rice OsDWARF4 and A. thaliana DWARF4 (47.4%, 37.7% and 41.7% amino-acid identities, respectively; Fig. 1a). In wild-type rice, OsDWARF4L1/D11 transcripts were most abundant in vegetative shoot apices, leaf sheaths and roots, and found at more moderate levels in other organs (Fig. 1d). They were decreased by exogenous brassinolide treatment, and increased in d61-3 and brd1-1 (Fig. 1e). These results indicate that OsDWARF4L1/D11 expression is regulated by a feedback mechanism, as is that of OsDWARF4.

----Phenotypes of rice mutants defective in brassinosteroid biosynthesis

In osdwarf4-1, Tos17 insertion into exon 6 causes a premature termination in domain C (arrowhead in Fig. 1b,c). Because a premature stop codon before the heme-binding domain caused loss-of-function of A. thaliana DWARF4 (ref. [7]), we consider osdwarf4-1 to be a null allele. Unexpectedly, osdwarf4-1 plants show only weak morphological phenotypes, such as a slightly dwarfed stature and more erect leaves (Fig. 2a,b). Both of these phenotypes were complemented by the introduction of an entire OsDWARF4 gene (data not shown). These results seem inconsistent with the fact that rice d61-3 and brd1-1, which are loss-of-function mutants of OsBRI1 and OsDWARF respectively, display severe dwarfing and have malformed leaves (for example, see brd1-1 in Fig. 2a), and suggest functional redundancy in brassinosteroid biosynthesis in rice.
Because null alleles of d11, such as d11-4, also show a weak phenotype[9] (Fig. 2a),OsDWARF4L1/D11 and OsDWARF4 may function redundantly. To confirm this possibility, researcher produced double mutants by crossing homozygous osdwarf4-1 and d11-4 plants. Among the F2 population, 34 of 576 plants showed severe dwarfing and malformed leaves with tortuous leaf blades, indistinguishable from the brassinosteroid-deficient brd1-1 phenotype (Fig. 2a,c,d). This ratio fits the theoretical 9:3:3:1 (wild-type:osdwarf4-1:d11-4:osdwarf4-1/d11-4) segregation predicted by mendelian inheritance. PCR and sequence analyses confirmed that these plants were osdwarf4-1/d11-4 double mutants (data not shown).
Endogenous levels of brassinosteroid intermediates in brd1 clearly indicate that OsDWARF catalyzes C-6 oxidation[4]. However, quantification in d2 could not determine the step(s) catalyzed by CYP90D2/D2, because CYP90D2/D2 functions redundantly with another gene, CYP90D3, and loss-of-function of CYP90D2/D2 have caused only limited defects in brassinosteroid biosynthesis[5]. Similarly, endogenous levels of brassinosteroid intermediates were obviously changed in the osdwarf4-1/d11-4 double mutant relative to osdwarf4-1 or d11 single mutants. Although brassinolide was not detected in either the wild-type or osdwarf4-1/d11-4, another bioactive brassinosteroid, castasterone (CS), was detected in the wild-type but not in osdwarf4-1/d11-4, confirming that osdwarf4-1/d11-4 is brassinosteroid-deficient (Fig. 2e). Levels of the other five intermediates downstream of C-22 hydroxylation—6-deoxocathasterone (6-DeoxoCT), 6-deoxoteasterone (6-DeoxoTE), 3-dehydro-6-deoxoteasterone (6-Deoxo3DT), 6-deoxotyphasterol (6-DeoxoTY) and 6-deoxocastasterone (6-DeoxoCS)—were also greatly reduced in osdwarf4-1/d11-4 (Fig. 2e). This indicates that both proteins are involved in C-22 hydroxylation. It is unclear why the upstream compound, campestanol, did not accumulate in osdwarf4-1/d11-4. Because A. thaliana CYP90B1/DWARF4 also catalyzes C-22 hydroxylation of other upstream compounds such as campesterol and (24R)-ergost-4-en-3-one[10], it is possible that campestanol is not the major substrate for CYP90B2/OsDWARF4 and CYP724B1/D11 in rice.

----Characterization of a field-grown osdwarf4-1 mutant

Because the harvest index (grain/(grain plus straw)) of rice is about 0.5, further increases in yield potential will have to involve increases in crop biomass driven by more net photosynthesis[11]. In rice, the contribution of lower leaves to photosynthesis is significant, even though their photosynthetic capacity is considerably less than that of upper leaves[12]. Erect leaves allow greater penetration of light to lower leaves, thereby optimizing canopy photosynthesis[13]. Therefore, the most likely explanation for the increase in above-ground biomass in osdwarf4-1 plots under dense planting conditions (more than 1.2 times that in wild-type under conventional planting condition) is that the shade of the upper leaves is minimized, and the lower leaves receive more light to drive higher rates of photosynthesis (Fig. 3 and Supplementary Table 2 online). No difference was observed in the heading date (flowering time) and the grain-filling period (time to maturity) between wild-type and osdwarf4-1 plants (data not shown). In the wild-type, the above-ground biomass was increased in response to higher fertilizer application under conventional planting (22.2 plants m-2), but was significantly decreased under dense planting with doubled fertilizer application owing to reduced photosynthesis caused by lodging (Fig. 3a). In contrast, the above-ground biomass of osdwarf4-1 was significantly (P < 0.01) increased by dense planting at each fertilizer rate. Consequently, the above-ground biomass in osdwarf4-1 plots under dense planting with standard and a 1.5-fold increase in fertilizer application was approx1.34 and 1.52 times that in wild-type plots under conventional cultivation conditions. The increased biomass depended mainly on accelerated formation of tillers, which have the ability to generate a panicle (panicle number, Fig. 3b). Both total and fertile grain numbers tended to increase in densely planted osdwarf4-1 plots (Fig. 3c), resulting in significantly increased grain yield even when the fertilizer application was not increased (for example, the osdwarf4-1 yield was 6.19 t ha-1 under dense planting with standard fertilizer application whereas the wild-type yield was 4.69 t ha-1 under conventional conditions; Fig. 3d). However, the increase in grain yield was not as much as that in above-ground biomass, because the harvest index of osdwarf4-1 (0.41) was slightly lower than that of the wild-type (0.44). A similar field test gave comparable results.

----Overexpression of OsDWARF4

Overexpression of DWARF in transgenic tomato increases plant height[14], and overexpression of DWARF4 in transgenic A. thaliana and tobacco increases vegetative growth and seed yield[15]. In rice, however, the role of brassinosteroid in regulating leaf angle has been known for over 40 years[16, 17]. Whereas brassinosteroid deficiency increased the erectness of leaves, the opposite phenotype (increased leaf angle) was observed in transgenic rice overexpressing OsDWARF4 (27.6° versus 10.0° in the wild-type, P < 0.01), and was inherited in their progeny (data not shown). Because inclined leaves are unfavorable for photosynthesis when plants are grown in a group, we have to carefully evaluate whether the overexpression of a brassinosteroid-biosynthesis gene in transgenic rice increases grain yield as is the case in transgenic plants overexpressing DWARF4.

===Conclusion===

More erect leaves improve not only light distribution but also the size and use of leaf nitrogen reservoirs for grain growth[13]. The erect leaf phenotype caused by brassinosteroid deficiency improved biomass production under field conditions, resulting in increased grain yield without the negative environmental effects associated with fertilizers. Recently identified rice homologs of A. thaliana BAS1 that encodes the brassinosteroid catabolic enzyme, CYP734A1 (ref. [18]). Overexpression of these rice homologs in transgenic rice successfully induced the brassinosteroid-deficient phenotypes (unpublished results). Although ideally, expression of these genes should be restricted to particular tissues and organs such as leaf laminae, it should be possible to generate erect leaf varieties by the introduction of a single transgene. In conclusion, engineering more erect leaves modulating brassinosteroid metabolism can be combined with dense planting to improve rice grain yields.

==Labs working on this gene==
Please input related labs here.

==References==
[1] Sakamoto, T. & Matsuoka, M. Generating high-yielding varieties by genetic manipulation of plant architecture. Curr. Opin. Biotech. 15, 144–147 (2004).
[2] Li, J. & Chory, J. A putative leucine-rich repeat receptor kinase involved in brassinosteroid signal transduction. Cell 90, 929–938 (1997).
[3] Yamamuro, C. et al. Loss of function of a rice brassinosteroid insensitive1 homolog prevents internode elongation and bending of the lamina joint. Plant Cell 12, 1591–1606 (2000).
[4] Hong, Z. et al. Loss-of-function of a rice brassinosteroid biosynthetic enzyme, C-6 oxidase, prevents the organized arrangement and polar elongation of cells in the leaves and stem. Plant J. 32, 495–508 (2002).
[5] Hong, Z. et al. A rice brassinosteroid-deficient mutant, ebisu dwarf (d2), is caused by a loss of function of a new member of cytochrome P450. Plant Cell 15, 2900–2910 (2003).
[6]Sakamoto T;Morinaka Y;Ohnishi T;Sunohara H;Fujioka S;Ueguchi Tanaka M;Mizutani M;Sakata K;Takatsuto S;Yoshida S;Tanaka H;Kitano H;Matsuoka M
Erect leaves caused by brassinosteroid deficiency increase biomass production and grain yield in rice
Nature Biotechnology, 2006, 24(1): 105-109.
[7] Choe, S. et al. The DWF4 gene of Arabidopsis encodes a cytochrome P450 that mediates multiple 22alpha-hydroxylation steps in brassinosteroid biosynthesis. Plant Cell 10, 231–243 (1998).
[8] Bancos, S. et al. Regulation of transcript levels of the Arabidopsis cytochrome P450 genes involved in brassinosteroid biosynthesis. Plant Physiol. 130, 504–513 (2002).
[9] Tanabe, S. et al. A novel cytochrome P450 is implicated in brassinosteroid biosynthesis via the characterization of a rice dwarf mutant, dwarf11, with reduced seed length. Plant Cell 17, 776–790 (2005).
[10] Fujioka, S., Takatsuto, S. & Yoshida, S. An early C-22 oxidation branch in brassinosteroid biosynthetic pathway. Plant Physiol. 130, 930–939 (2002).
[11] Mann, C.C. Crop scientists seek a new revolution. Science 283, 310–314 (1999).
[12] Horton, P. Prospects for crop improvement through the genetic manipulation of photosynthesis: morphological and biochemical aspects of light capture. J. Exp. Bot. 51, 475–485 (2000).
[13] Sinclair, T.R. & Sheehy, J.E. Erect leaves and photosynthesis in rice. Science 283, 1455 (1999).
[14] Bishop, G.J. et al. The tomato DWARF enzyme catalyses C-6 oxidation in brassinosteroid biosynthesis. Proc. Natl. Acad. Sci. USA 96, 1761–1766 (1999).
[15] Choe, S. et al. Overexpression of DWARF4 in the brassinosteroid biosynthetic pathway results in increased vegetative growth and seed yield in Arabidopsis. Plant J. 26, 573–582 (2001).
[16] Maeda, E. Rate of lamina inclination in excised rice leaves. Physiol. Plant. 18, 813–827 (1965).
[17] Wada, K. et al. Brassinolide and homobrassinolide promotion of lamina inclination of rice seedlings. Plant Cell Physiol. 22, 323–325 (1981).
[18] Neff, M.M. et al. BAS1: A gene regulating brassinosteroid levels and light responsiveness in Arabidopsis. Proc. Natl. Acad. Sci. USA 96, 15316–15323 (1999).

==Structured Information==
{{JaponicaGene|
GeneName = Os03g0227700|
Description = Cytochrome P450 family protein|
Version = NM_001055982.2 GI:297600576 GeneID:4332134|
Length = 6529 bp|
Definition = Oryza sativa Japonica Group Os03g0227700, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 3|Chromosome 3]]|
AP = Chromosome 3:6765109..6771637|
CDS = 6765109..6765374,6765543..6765867,6765971..6766123,6768937..6769188,6769337..6769429 ,6770222..6770300,6770398..6770504,6771392..6771637|
GCID = <gbrowseImage1>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</cdnaseq>|
AA = <aaseq>MAAMMASITSELLFFLPFILLALLTFYTTTVAKCHGGHWWRGGT TPAKRKRMNLPPGAAGWPLVGETFGYLRAHPATSVGRFMEQHIARYGKIYRSSLFGER TVVSADAGLNRYILQNEGRLFECSYPRSIGGILGKWSMLVLVGDPHREMRAISLNFLS SVRLRAVLLPEVERHTLLVLRAWPPSSTFSAQHQAKKFTFNLMAKNIMSMDPGEEETE RLRREYITFMKGVVSAPLNLPGTPYWKALKSRAAILGVIERKMEERVEKLSKEDASVE QDDLLGWALKQSNLSKEQILDLLLSLLFAGHETSSMALALAIFFLEGCPKAVQELREE HLGIARRQRLRGECKLSWEDYKEMVFTQCVINETLRLGNVVRFLHRKVIKDVHYKGYD IPSGWKILPVLAAVHLDSSLYEDPQRFNPWRWKSSGSSGGLAQSSSFMPYGGGTRLCA GSELAKLEMAVFLHHLVLNFRWELAEPDQAFVFPFVDFPKGLPIRVHRIAQDDEQE</aaseq>|
DNA = <dnaseqindica>1..266#435..759#863..1015#3829..4080#4229..4321#5114..5192#5290..5396#6284..6529#atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtcggtggtttcattactcggtacacgcacacagctatataagcacgcgcactgctcgcctcgccgtgaagcaagcttagctagtagctaggatatcgatcgatcggtaataataggctgacggcgagctggggggtggcgtgcggttgcgtgcgtgcgtgcatgcaggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaaggtacaacacacatcgtcctccattaagttgagcatgcatatatcaatcagctcgctctagcgatcaatatcaatgaaacgcgcgtttgcgtgcgtgcgtgcagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaaggtacgtactcgtagctactagctactccctccgtttcgaaatgtttgacgccgttgactttttatcacatatttgaccgttcgttttatttaaaaaaattaagtaattattaattattttcctatcatttgattcattgttaaatatatttttatgtaggcatataattttacatatctcacaaaagtttttaaataagacgaacggtcaaacatttgctaaaaagtcaacggtgtcaaatatttcgaaacggaggaagtagctagctaccaccccacccgtgcggattgatcgatcgatcgatgtccctctctgcctacacacaccctgcgccccggccgtacgcgcgcaaagtcgtgcgcgcatgagttttactccttccgtttcaaaaaaaaaacgacctagacaagtaatatgatgtaacctagaacaacgaatctatacaggagtatatccaaattcattgtcctagattacatcatatctttgtctatgttgccttttttaggaggagagagtactttacatatgagttatgaccggccggccaggttcctttcttcccctattccgtatgcatactataagcagctgcatataggcatatactactatacccagcttagctaaagcaaacaacttgcaacgacgatcatgtgcaggtccctgttgacagaggttagagatgtaaaccatttttgttgttttcattatctaaagagaaaacaagttactacgacgatcgagatgtacgcaaggggatcatatttatattggagaggagagagagatgatttgcgacttgtgagtgataagtgaatatatatactatatatttgtgtaggagtgacgatgacttccctgtcaatgttttgtttggtatgcacacggggggaaaagagtttagagaaaggaattggctagagagggccaggtttagtttctttaatttttgtttggtggttttgtgtgtagctagtccgcatgcatgatatatcatatatatgcatggcccatcattagatatatatacatgacggcagtgcacttattatatatatgtaggggcatatacaaggtgtatgtgtttagaattgtggatggtccaggaattgttgcagctagttctatactagcatagtgaattagtgatccatatcgagatcaggtccaatcggtcagctcgcacagcgggtcttttagtacgatttgttcggacaaattatcttgcctttgagataatcaacataggattgaaactgcatactagcaatagaccatagctatttggcaatatagaaagctaaaaaatgttgattgaaaggtgcgtgcatgcatgctagcagtatgtcttttaggccaggtccagtttttttttttttctcaaaggcagttgcttaaggtgaaattctagcagaggtcatatgaatatatttattatcttccaaacatgagccaaaacctatatatagttaactcaaaaaaccatgcattggtacttcttctgagtatgcatcctggggactgatcgatcatgcagattcttttctttctttttccctggggacgtactgatcatcactcatgtgagcataaaatgttgcctactttttactgcatgtgaggtcacactcaaatgatctttattgattgcctcagcatgcatgctgctagctttcacgatctctctccgatacacacaccctgtgctgtgctcaacagcatgcaacctggcaggtggaatttttaatctgttggaaagtaaatgaccaaccttcataatttggcatgattatatgctggggaatctatcatttatcatgattcaaccgctgaccaaactctttgccacacacccgaaccacattaatttcgttaagcggatgattagccagcaaccttcacagtttggttttaattagttaatgaaagggtgcttgtagttttattagctgaagagacactgcagtactcctactgagcgttctccttaattgggaaatagatctctgcagtactatcctttaactttaagtgaataacatgtagctaggtctcacacttatcaggcccacacatcagtgatccaagtcaaatggcaatacaccataggagattcttccactaattggctaacaaaaataaactgtgatcactgatcacagctgctaacactataacacattaatttgcatgtctcatgaaatgccagatgagttggattaattatctatcttatgttcagtgccgaggaatatacaaggaaacatcattggtatagttgtggagttctgagtgcattatctcacccagtacggtcaaattaatcctagtctccgtaaatatggttgtgtgcactcttagttaattagtgtagacccggaaattatactttgtcgagaatgttgttatccgtgtgttaagtttcaagtaccctgcaggcctataactaaacactgccaacatggatgtttaagaatgaattatacatacagcgcacttggaaaagaaaaggggtagctacccagctgaagaacacctctctctcatttgaaggtcgacacaatatgtgtcttttctatatcgaaaggaaacgacattttttctcttcaaaacagaaagctatgacaagagacataatcctagactagtaggcggttaggcaacaccatatctgaatttgataggtgtcccacatattttttttcttttcctgataagaaggaacttttcaccgtgagcttatttttttctgttctatgtcgacattagttccatgatgtttactgtgtatttttactgttcccctgcagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggtacagagaattacagattcgaaattaacatctactgtcactagctcagtgctacgtttaactgtagagcaacaattgcagaaagcgaaggcatcagaaatgacatgggaatcgattagttacactcacactgacgcgtgacatgcaggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtaaggccccccctactttgttccttccgatccctcctccctaaacgctactaatataaatcggtgcaaaggctttgtgctagcgtagctcttgctgttttgtttggtcacttgcgtagagagatagcaacttcccgaaatgtcatgtacgaatggagactgaaagctctgcgaaataattgccgggccacgtactacagtacttacgctccctctcttcagtttggatcgatcgagctaagaaggcgcacgcatgtgtgcattgatcgatagagtcgatctgttctgccttgtaagctagccagctgaggttctttcgccttggaagtttctacagggtgaggcccatcgtcttgcaacgatccaaatgccatgacaagtacctgttgttgtaggcttccagtagcacggcctgcccccaagaaggtatctgaatcttctgcattgctgttcagacggcagagcagaacatgccgaaataccattccagtcagttgcagtatatgtgtgtggtggtatcagataaaggagcagagtaatttgaggccttagaaattcagatgagttgctcgacaattgtcgcctaccgatcgagaataagttgcatgcatgtatgtgatgtaattatgggtcttagtcagcaggatagtttagtcgctgcagtactgtagattgtagtacttgcgttcaaagcagacccaaaacagaacttgcattcaggaaacaaaacatgaaaaaagaaatatacagtacttggcactgatatttcttttgtttgggtccttattgcaggtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggtgagtagctcgatttcactcttcttccctgaatcgacctctcctctatctgacagcagaagctaactgaactgaactgaactgaactgaactgaaggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaaggtcagtcgcagtaggattatcagtgaatctaatctattaatctcggtttaattgctgggattcaaggaaaagaaaatactgggacaagcgcggatgtcgctgttgggcagtgtctgtcaatcataagcgaaggggatcatgtcagaatctgcaaaagcagcacaggctaaacgcgtgcttgcctcgagcagcaatcatgcagagaggcaagctccgcgataggaacatgcatggtgaatcactgaacctgatatgaaagtgaatgctcagctcagctcgctcgctcgcgcgcgcatgtgctttctcatttcgccgtgttgcgtgctagctagtcaatggtgttcccacgaattgttttctcttctcatcacacaaagtcagtggtgaaaaccgaggtgcatgtgcggtaatggaacgggggattaggcggcgacgccgtgatggattatgcattcgtgcgagaaaaatccaggagcgttggtccaaatcgcaagcggatcaaaccgaaccaatcagtggacgattggagccctgcagtccatccatcacatctccattcttatccgctgcgactcacggacattagttgttgaggggtgaacgacgcatgtgtgtatgtatgctcgatcgatcgatcgaacaatggatctgtgtgtggtgtagttgccaacgccgacgagattatctgctcgtctgcatgcagccgcgaaaaagacatgtgtccggattatcctgcaggagattgagcaagataatctgctgcgcaaaaactgcagccaatcgatcattccatgacgcgaaaaaaaaaacaataattactactgctgatacgataacttcgcttcgaattgttgacacacaagtgttcttcttttgttcgattgcagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001055982.2 RefSeq:Os03g0227700]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 3]]
[[Category:Chromosome 3]]

Os03g0227700

2014-05-24T13:34:03Z

LQ: /* Annotated Information */

Please input one-sentence summary here.
''OsDWAR''F4 is expressed in young elongating tissues.
==Annotated Information==

Brassinosteroids influence both plant height and leaf erectness in rice[1]. Arabidopsis thaliana BRASSINOSTEROID INSENSITIVE1 (bri1) mutants, which are deficient in the brassinosteroid receptor, have a severe dwarf phenotype[2]. The rice mutants d61-1 and d61-2, which are weak mutant alleles of OsBRI1, a homolog of A. thaliana BRI1, have both a semi-dwarf phenotype and more erect leaves[3]. In rice, an erect leaf phenotype has also been associated with loss-of-function, brassinosteroid-deficient mutants such as brassinosteroid-deficient dwarf1 (brd1), which is defective in OsDWARF (the homolog of tomato CYP85A1/DWARF[4]), and ebisu dwarf (d2), which is defective in CYP90D2/D2 (ref. [5]). However, the grain yield of d61, brd1 and d2 mutants is decreased because of morphological alterations in their reproductive development.

===Function===

New cultivars with very erect leaves, which increase light capture for photosynthesis and nitrogen storage for grain filling, may have increased grain yields.The erect leaf phenotype of a rice brassinosteroid-deficient mutant, osdwarf4-1, is associated with enhanced grain yields under conditions of dense planting, even without extra fertilizer. Molecular and biochemical studies reveal that two different cytochrome P450s, CYP90B2/OsDWARF4 and CYP724B1/D11, function redundantly in C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis. Therefore, despite the central role of brassinosteroids in plant growth and development, mutation of OsDWARF4 alone causes only limited defects in brassinosteroid biosynthesis and plant morphology[6].

===Expression===
The affected gene, OsDWARF4, was identified by characterizing the site of insertion of the Tos17 retrotransposon.

----Molecular characterization of OsDWARF4 and OsDWARF4L1/D11

OsDWARF4 is a homolog of A. thaliana DWARF4, which encodes a cytochrome P450, CYP90B1 (Fig. 1a). CYP90B1/DWARF4 catalyzes C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis[7]. Only one DWARF4 homolog was identified in the rice genome. OsDWARF4 is located on the short arm of chromosome 3 (35 cM). Its predicted open reading frame consists of eight exons, and encodes a protein of 502 amino acids (Fig. 1b,c). OsDWARF4 is most closely related to DWARF4 (66.3% amino-acid identity; Fig. 1a). The structure of OsDWARF4 is similar to that of DWARF4 throughout its length: five domains found in cytochrome P450s—proline rich, A (also referred to as dioxygen binding), B (steroid binding), C and heme binding—are highly conserved (Fig. 1c). In wild-type rice, OsDWARF4 transcripts are most abundant in leaf blades and roots, but are also found in all other plant parts tested (Fig. 1d). Transcript levels were decreased by exogenous brassinolide treatment, and increased in brassinosteroid-insensitive d61-3 and brassinosteroid-deficient brd1-1 (Fig. 1e). These results suggest that OsDWARF4 expression is feedback regulated, as is that of A. thaliana brassinosteroid-biosynthesis genes[8].
OsDWARF4L1 was identified as an OsDWARF4-like gene by in silico screening of rice DNA databases. OsDWARF4L1 is located on the long arm of chromosome 4 (70.9 cM). A recent study demonstrated that a rice semi-dwarf and short-grain mutant, d11, has a loss-of-function mutation in the OsDWARF4L1 gene[9]. OsDWARF4L1/D11 shows the highest homology with A. thaliana CYP724A1, and is closely related to rice OsDWARF4 and A. thaliana DWARF4 (47.4%, 37.7% and 41.7% amino-acid identities, respectively; Fig. 1a). In wild-type rice, OsDWARF4L1/D11 transcripts were most abundant in vegetative shoot apices, leaf sheaths and roots, and found at more moderate levels in other organs (Fig. 1d). They were decreased by exogenous brassinolide treatment, and increased in d61-3 and brd1-1 (Fig. 1e). These results indicate that OsDWARF4L1/D11 expression is regulated by a feedback mechanism, as is that of OsDWARF4.

----Phenotypes of rice mutants defective in brassinosteroid biosynthesis

In osdwarf4-1, Tos17 insertion into exon 6 causes a premature termination in domain C (arrowhead in Fig. 1b,c). Because a premature stop codon before the heme-binding domain caused loss-of-function of A. thaliana DWARF4 (ref. [7]), we consider osdwarf4-1 to be a null allele. Unexpectedly, osdwarf4-1 plants show only weak morphological phenotypes, such as a slightly dwarfed stature and more erect leaves (Fig. 2a,b). Both of these phenotypes were complemented by the introduction of an entire OsDWARF4 gene (data not shown). These results seem inconsistent with the fact that rice d61-3 and brd1-1, which are loss-of-function mutants of OsBRI1 and OsDWARF respectively, display severe dwarfing and have malformed leaves (for example, see brd1-1 in Fig. 2a), and suggest functional redundancy in brassinosteroid biosynthesis in rice.
Because null alleles of d11, such as d11-4, also show a weak phenotype[9] (Fig. 2a),OsDWARF4L1/D11 and OsDWARF4 may function redundantly. To confirm this possibility, researcher produced double mutants by crossing homozygous osdwarf4-1 and d11-4 plants. Among the F2 population, 34 of 576 plants showed severe dwarfing and malformed leaves with tortuous leaf blades, indistinguishable from the brassinosteroid-deficient brd1-1 phenotype (Fig. 2a,c,d). This ratio fits the theoretical 9:3:3:1 (wild-type:osdwarf4-1:d11-4:osdwarf4-1/d11-4) segregation predicted by mendelian inheritance. PCR and sequence analyses confirmed that these plants were osdwarf4-1/d11-4 double mutants (data not shown).
Endogenous levels of brassinosteroid intermediates in brd1 clearly indicate that OsDWARF catalyzes C-6 oxidation[4]. However, quantification in d2 could not determine the step(s) catalyzed by CYP90D2/D2, because CYP90D2/D2 functions redundantly with another gene, CYP90D3, and loss-of-function of CYP90D2/D2 have caused only limited defects in brassinosteroid biosynthesis[5]. Similarly, endogenous levels of brassinosteroid intermediates were obviously changed in the osdwarf4-1/d11-4 double mutant relative to osdwarf4-1 or d11 single mutants. Although brassinolide was not detected in either the wild-type or osdwarf4-1/d11-4, another bioactive brassinosteroid, castasterone (CS), was detected in the wild-type but not in osdwarf4-1/d11-4, confirming that osdwarf4-1/d11-4 is brassinosteroid-deficient (Fig. 2e). Levels of the other five intermediates downstream of C-22 hydroxylation—6-deoxocathasterone (6-DeoxoCT), 6-deoxoteasterone (6-DeoxoTE), 3-dehydro-6-deoxoteasterone (6-Deoxo3DT), 6-deoxotyphasterol (6-DeoxoTY) and 6-deoxocastasterone (6-DeoxoCS)—were also greatly reduced in osdwarf4-1/d11-4 (Fig. 2e). This indicates that both proteins are involved in C-22 hydroxylation. It is unclear why the upstream compound, campestanol, did not accumulate in osdwarf4-1/d11-4. Because A. thaliana CYP90B1/DWARF4 also catalyzes C-22 hydroxylation of other upstream compounds such as campesterol and (24R)-ergost-4-en-3-one[10], it is possible that campestanol is not the major substrate for CYP90B2/OsDWARF4 and CYP724B1/D11 in rice.

----Characterization of a field-grown osdwarf4-1 mutant

Because the harvest index (grain/(grain plus straw)) of rice is about 0.5, further increases in yield potential will have to involve increases in crop biomass driven by more net photosynthesis[11]. In rice, the contribution of lower leaves to photosynthesis is significant, even though their photosynthetic capacity is considerably less than that of upper leaves[12]. Erect leaves allow greater penetration of light to lower leaves, thereby optimizing canopy photosynthesis[13]. Therefore, the most likely explanation for the increase in above-ground biomass in osdwarf4-1 plots under dense planting conditions (more than 1.2 times that in wild-type under conventional planting condition) is that the shade of the upper leaves is minimized, and the lower leaves receive more light to drive higher rates of photosynthesis (Fig. 3 and Supplementary Table 2 online). No difference was observed in the heading date (flowering time) and the grain-filling period (time to maturity) between wild-type and osdwarf4-1 plants (data not shown). In the wild-type, the above-ground biomass was increased in response to higher fertilizer application under conventional planting (22.2 plants m-2), but was significantly decreased under dense planting with doubled fertilizer application owing to reduced photosynthesis caused by lodging (Fig. 3a). In contrast, the above-ground biomass of osdwarf4-1 was significantly (P < 0.01) increased by dense planting at each fertilizer rate. Consequently, the above-ground biomass in osdwarf4-1 plots under dense planting with standard and a 1.5-fold increase in fertilizer application was approx1.34 and 1.52 times that in wild-type plots under conventional cultivation conditions. The increased biomass depended mainly on accelerated formation of tillers, which have the ability to generate a panicle (panicle number, Fig. 3b). Both total and fertile grain numbers tended to increase in densely planted osdwarf4-1 plots (Fig. 3c), resulting in significantly increased grain yield even when the fertilizer application was not increased (for example, the osdwarf4-1 yield was 6.19 t ha-1 under dense planting with standard fertilizer application whereas the wild-type yield was 4.69 t ha-1 under conventional conditions; Fig. 3d). However, the increase in grain yield was not as much as that in above-ground biomass, because the harvest index of osdwarf4-1 (0.41) was slightly lower than that of the wild-type (0.44). A similar field test gave comparable results.

----Overexpression of OsDWARF4

Overexpression of DWARF in transgenic tomato increases plant height[14], and overexpression of DWARF4 in transgenic A. thaliana and tobacco increases vegetative growth and seed yield[15]. In rice, however, the role of brassinosteroid in regulating leaf angle has been known for over 40 years[16, 17]. Whereas brassinosteroid deficiency increased the erectness of leaves, the opposite phenotype (increased leaf angle) was observed in transgenic rice overexpressing OsDWARF4 (27.6° versus 10.0° in the wild-type, P < 0.01), and was inherited in their progeny (data not shown). Because inclined leaves are unfavorable for photosynthesis when plants are grown in a group, we have to carefully evaluate whether the overexpression of a brassinosteroid-biosynthesis gene in transgenic rice increases grain yield as is the case in transgenic plants overexpressing DWARF4.

===Conclusion===

More erect leaves improve not only light distribution but also the size and use of leaf nitrogen reservoirs for grain growth[13]. The erect leaf phenotype caused by brassinosteroid deficiency improved biomass production under field conditions, resulting in increased grain yield without the negative environmental effects associated with fertilizers. Recently identified rice homologs of A. thaliana BAS1 that encodes the brassinosteroid catabolic enzyme, CYP734A1 (ref. [18]). Overexpression of these rice homologs in transgenic rice successfully induced the brassinosteroid-deficient phenotypes (unpublished results). Although ideally, expression of these genes should be restricted to particular tissues and organs such as leaf laminae, it should be possible to generate erect leaf varieties by the introduction of a single transgene. In conclusion, engineering more erect leaves modulating brassinosteroid metabolism can be combined with dense planting to improve rice grain yields.

==Labs working on this gene==
Please input related labs here.

==References==
[1]Sakamoto T;Morinaka Y;Ohnishi T;Sunohara H;Fujioka S;Ueguchi Tanaka M;Mizutani M;Sakata K;Takatsuto S;Yoshida S;Tanaka H;Kitano H;Matsuoka M
Erect leaves caused by brassinosteroid deficiency increase biomass production and grain yield in rice
Nature Biotechnology, 2006, 24(1): 105-109.
[1] Sinclair, T.R. & Sheehy, J.E. Erect leaves and photosynthesis in rice. Science 283, 1455 (1999).
[2] Evans, L.T. Crop Evolution, Adaptation and Yield (Cambridge Univ. Press, Cambridge, 1993).
[3] Peng, J. et al. 'Green revolution' genes encode mutant gibberellin response modulators. Nature 400, 256–261 (1999).
[4] Sasaki, A. et al. Green revolution: a mutant gibberellin-synthesis gene in rice. Nature 416, 701–702 (2002).
[5] Sakamoto, T. et al. Genetic manipulation of gibberellin metabolism in transgenic rice. Nat. Biotechnol. 21, 909–913 (2003).
[6] Sakamoto, T. & Matsuoka, M. Generating high-yielding varieties by genetic manipulation of plant architecture. Curr. Opin. Biotech. 15, 144–147 (2004).
[7] Li, J. & Chory, J. A putative leucine-rich repeat receptor kinase involved in brassinosteroid signal transduction. Cell 90, 929–938 (1997).
[8] Yamamuro, C. et al. Loss of function of a rice brassinosteroid insensitive1 homolog prevents internode elongation and bending of the lamina joint. Plant Cell 12, 1591–1606 (2000).
[9] Hong, Z. et al. Loss-of-function of a rice brassinosteroid biosynthetic enzyme, C-6 oxidase, prevents the organized arrangement and polar elongation of cells in the leaves and stem. Plant J. 32, 495–508 (2002).
[10] Hong, Z. et al. A rice brassinosteroid-deficient mutant, ebisu dwarf (d2), is caused by a loss of function of a new member of cytochrome P450. Plant Cell 15, 2900–2910 (2003).
[11] Choe, S. et al. The DWF4 gene of Arabidopsis encodes a cytochrome P450 that mediates multiple 22alpha-hydroxylation steps in brassinosteroid biosynthesis. Plant Cell 10, 231–243 (1998).
[12] Bancos, S. et al. Regulation of transcript levels of the Arabidopsis cytochrome P450 genes involved in brassinosteroid biosynthesis. Plant Physiol. 130, 504–513 (2002).
[13] Tanabe, S. et al. A novel cytochrome P450 is implicated in brassinosteroid biosynthesis via the characterization of a rice dwarf mutant, dwarf11, with reduced seed length. Plant Cell 17, 776–790 (2005).
[14] Fujioka, S., Takatsuto, S. & Yoshida, S. An early C-22 oxidation branch in brassinosteroid biosynthetic pathway. Plant Physiol. 130, 930–939 (2002).
[15] Mann, C.C. Crop scientists seek a new revolution. Science 283, 310–314 (1999).
[16] Horton, P. Prospects for crop improvement through the genetic manipulation of photosynthesis: morphological and biochemical aspects of light capture. J. Exp. Bot. 51, 475–485 (2000).
[17] Bishop, G.J. et al. The tomato DWARF enzyme catalyses C-6 oxidation in brassinosteroid biosynthesis. Proc. Natl. Acad. Sci. USA 96, 1761–1766 (1999).
[18] Choe, S. et al. Overexpression of DWARF4 in the brassinosteroid biosynthetic pathway results in increased vegetative growth and seed yield in Arabidopsis. Plant J. 26, 573–582 (2001).
[19] Maeda, E. Rate of lamina inclination in excised rice leaves. Physiol. Plant. 18, 813–827 (1965).
[20] Wada, K. et al. Brassinolide and homobrassinolide promotion of lamina inclination of rice seedlings. Plant Cell Physiol. 22, 323–325 (1981).
[21] Neff, M.M. et al. BAS1: A gene regulating brassinosteroid levels and light responsiveness in Arabidopsis. Proc. Natl. Acad. Sci. USA 96, 15316–15323 (1999).
[22] Sakamoto, T. et al. An overview of gibberellin metabolism enzyme genes and their related mutants in rice. Plant Physiol. 134, 1642–1653 (2004).
[23] Saito, S. et al. Arabidopsis CYP707As encode (+)-abscisic acid 8'-hydroxylase, a key enzyme in the oxidative catabolism of abscisic acid. Plant Physiol. 134, 1439–1449 (2004).
[24] Fujita, S. et al. Arabidopsis CYP90B1 catalyzes the early C-22 hydroxylation of C27, C28, and C29 sterols. Plant J. 45, in press.
[25] Hiei, Y., Ohta, S., Komari, T. & Kumashiro, T. Efficient transformation of rice (Oryza sativa L.) mediated by Agrobacterium and sequence analysis of boundaries of the T-DNA. Plant J. 6, 271–282 (1994).

==Structured Information==
{{JaponicaGene|
GeneName = Os03g0227700|
Description = Cytochrome P450 family protein|
Version = NM_001055982.2 GI:297600576 GeneID:4332134|
Length = 6529 bp|
Definition = Oryza sativa Japonica Group Os03g0227700, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 3|Chromosome 3]]|
AP = Chromosome 3:6765109..6771637|
CDS = 6765109..6765374,6765543..6765867,6765971..6766123,6768937..6769188,6769337..6769429 ,6770222..6770300,6770398..6770504,6771392..6771637|
GCID = <gbrowseImage1>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</cdnaseq>|
AA = <aaseq>MAAMMASITSELLFFLPFILLALLTFYTTTVAKCHGGHWWRGGT TPAKRKRMNLPPGAAGWPLVGETFGYLRAHPATSVGRFMEQHIARYGKIYRSSLFGER TVVSADAGLNRYILQNEGRLFECSYPRSIGGILGKWSMLVLVGDPHREMRAISLNFLS SVRLRAVLLPEVERHTLLVLRAWPPSSTFSAQHQAKKFTFNLMAKNIMSMDPGEEETE RLRREYITFMKGVVSAPLNLPGTPYWKALKSRAAILGVIERKMEERVEKLSKEDASVE QDDLLGWALKQSNLSKEQILDLLLSLLFAGHETSSMALALAIFFLEGCPKAVQELREE HLGIARRQRLRGECKLSWEDYKEMVFTQCVINETLRLGNVVRFLHRKVIKDVHYKGYD IPSGWKILPVLAAVHLDSSLYEDPQRFNPWRWKSSGSSGGLAQSSSFMPYGGGTRLCA GSELAKLEMAVFLHHLVLNFRWELAEPDQAFVFPFVDFPKGLPIRVHRIAQDDEQE</aaseq>|
DNA = <dnaseqindica>1..266#435..759#863..1015#3829..4080#4229..4321#5114..5192#5290..5396#6284..6529#atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtcggtggtttcattactcggtacacgcacacagctatataagcacgcgcactgctcgcctcgccgtgaagcaagcttagctagtagctaggatatcgatcgatcggtaataataggctgacggcgagctggggggtggcgtgcggttgcgtgcgtgcgtgcatgcaggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaaggtacaacacacatcgtcctccattaagttgagcatgcatatatcaatcagctcgctctagcgatcaatatcaatgaaacgcgcgtttgcgtgcgtgcgtgcagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaaggtacgtactcgtagctactagctactccctccgtttcgaaatgtttgacgccgttgactttttatcacatatttgaccgttcgttttatttaaaaaaattaagtaattattaattattttcctatcatttgattcattgttaaatatatttttatgtaggcatataattttacatatctcacaaaagtttttaaataagacgaacggtcaaacatttgctaaaaagtcaacggtgtcaaatatttcgaaacggaggaagtagctagctaccaccccacccgtgcggattgatcgatcgatcgatgtccctctctgcctacacacaccctgcgccccggccgtacgcgcgcaaagtcgtgcgcgcatgagttttactccttccgtttcaaaaaaaaaacgacctagacaagtaatatgatgtaacctagaacaacgaatctatacaggagtatatccaaattcattgtcctagattacatcatatctttgtctatgttgccttttttaggaggagagagtactttacatatgagttatgaccggccggccaggttcctttcttcccctattccgtatgcatactataagcagctgcatataggcatatactactatacccagcttagctaaagcaaacaacttgcaacgacgatcatgtgcaggtccctgttgacagaggttagagatgtaaaccatttttgttgttttcattatctaaagagaaaacaagttactacgacgatcgagatgtacgcaaggggatcatatttatattggagaggagagagagatgatttgcgacttgtgagtgataagtgaatatatatactatatatttgtgtaggagtgacgatgacttccctgtcaatgttttgtttggtatgcacacggggggaaaagagtttagagaaaggaattggctagagagggccaggtttagtttctttaatttttgtttggtggttttgtgtgtagctagtccgcatgcatgatatatcatatatatgcatggcccatcattagatatatatacatgacggcagtgcacttattatatatatgtaggggcatatacaaggtgtatgtgtttagaattgtggatggtccaggaattgttgcagctagttctatactagcatagtgaattagtgatccatatcgagatcaggtccaatcggtcagctcgcacagcgggtcttttagtacgatttgttcggacaaattatcttgcctttgagataatcaacataggattgaaactgcatactagcaatagaccatagctatttggcaatatagaaagctaaaaaatgttgattgaaaggtgcgtgcatgcatgctagcagtatgtcttttaggccaggtccagtttttttttttttctcaaaggcagttgcttaaggtgaaattctagcagaggtcatatgaatatatttattatcttccaaacatgagccaaaacctatatatagttaactcaaaaaaccatgcattggtacttcttctgagtatgcatcctggggactgatcgatcatgcagattcttttctttctttttccctggggacgtactgatcatcactcatgtgagcataaaatgttgcctactttttactgcatgtgaggtcacactcaaatgatctttattgattgcctcagcatgcatgctgctagctttcacgatctctctccgatacacacaccctgtgctgtgctcaacagcatgcaacctggcaggtggaatttttaatctgttggaaagtaaatgaccaaccttcataatttggcatgattatatgctggggaatctatcatttatcatgattcaaccgctgaccaaactctttgccacacacccgaaccacattaatttcgttaagcggatgattagccagcaaccttcacagtttggttttaattagttaatgaaagggtgcttgtagttttattagctgaagagacactgcagtactcctactgagcgttctccttaattgggaaatagatctctgcagtactatcctttaactttaagtgaataacatgtagctaggtctcacacttatcaggcccacacatcagtgatccaagtcaaatggcaatacaccataggagattcttccactaattggctaacaaaaataaactgtgatcactgatcacagctgctaacactataacacattaatttgcatgtctcatgaaatgccagatgagttggattaattatctatcttatgttcagtgccgaggaatatacaaggaaacatcattggtatagttgtggagttctgagtgcattatctcacccagtacggtcaaattaatcctagtctccgtaaatatggttgtgtgcactcttagttaattagtgtagacccggaaattatactttgtcgagaatgttgttatccgtgtgttaagtttcaagtaccctgcaggcctataactaaacactgccaacatggatgtttaagaatgaattatacatacagcgcacttggaaaagaaaaggggtagctacccagctgaagaacacctctctctcatttgaaggtcgacacaatatgtgtcttttctatatcgaaaggaaacgacattttttctcttcaaaacagaaagctatgacaagagacataatcctagactagtaggcggttaggcaacaccatatctgaatttgataggtgtcccacatattttttttcttttcctgataagaaggaacttttcaccgtgagcttatttttttctgttctatgtcgacattagttccatgatgtttactgtgtatttttactgttcccctgcagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggtacagagaattacagattcgaaattaacatctactgtcactagctcagtgctacgtttaactgtagagcaacaattgcagaaagcgaaggcatcagaaatgacatgggaatcgattagttacactcacactgacgcgtgacatgcaggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtaaggccccccctactttgttccttccgatccctcctccctaaacgctactaatataaatcggtgcaaaggctttgtgctagcgtagctcttgctgttttgtttggtcacttgcgtagagagatagcaacttcccgaaatgtcatgtacgaatggagactgaaagctctgcgaaataattgccgggccacgtactacagtacttacgctccctctcttcagtttggatcgatcgagctaagaaggcgcacgcatgtgtgcattgatcgatagagtcgatctgttctgccttgtaagctagccagctgaggttctttcgccttggaagtttctacagggtgaggcccatcgtcttgcaacgatccaaatgccatgacaagtacctgttgttgtaggcttccagtagcacggcctgcccccaagaaggtatctgaatcttctgcattgctgttcagacggcagagcagaacatgccgaaataccattccagtcagttgcagtatatgtgtgtggtggtatcagataaaggagcagagtaatttgaggccttagaaattcagatgagttgctcgacaattgtcgcctaccgatcgagaataagttgcatgcatgtatgtgatgtaattatgggtcttagtcagcaggatagtttagtcgctgcagtactgtagattgtagtacttgcgttcaaagcagacccaaaacagaacttgcattcaggaaacaaaacatgaaaaaagaaatatacagtacttggcactgatatttcttttgtttgggtccttattgcaggtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggtgagtagctcgatttcactcttcttccctgaatcgacctctcctctatctgacagcagaagctaactgaactgaactgaactgaactgaactgaaggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaaggtcagtcgcagtaggattatcagtgaatctaatctattaatctcggtttaattgctgggattcaaggaaaagaaaatactgggacaagcgcggatgtcgctgttgggcagtgtctgtcaatcataagcgaaggggatcatgtcagaatctgcaaaagcagcacaggctaaacgcgtgcttgcctcgagcagcaatcatgcagagaggcaagctccgcgataggaacatgcatggtgaatcactgaacctgatatgaaagtgaatgctcagctcagctcgctcgctcgcgcgcgcatgtgctttctcatttcgccgtgttgcgtgctagctagtcaatggtgttcccacgaattgttttctcttctcatcacacaaagtcagtggtgaaaaccgaggtgcatgtgcggtaatggaacgggggattaggcggcgacgccgtgatggattatgcattcgtgcgagaaaaatccaggagcgttggtccaaatcgcaagcggatcaaaccgaaccaatcagtggacgattggagccctgcagtccatccatcacatctccattcttatccgctgcgactcacggacattagttgttgaggggtgaacgacgcatgtgtgtatgtatgctcgatcgatcgatcgaacaatggatctgtgtgtggtgtagttgccaacgccgacgagattatctgctcgtctgcatgcagccgcgaaaaagacatgtgtccggattatcctgcaggagattgagcaagataatctgctgcgcaaaaactgcagccaatcgatcattccatgacgcgaaaaaaaaaacaataattactactgctgatacgataacttcgcttcgaattgttgacacacaagtgttcttcttttgttcgattgcagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001055982.2 RefSeq:Os03g0227700]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 3]]
[[Category:Chromosome 3]]

Os03g0227700

2014-05-24T13:33:12Z

LQ: /* Annotated Information */

Please input one-sentence summary here.
''OsDWAR''F4 is expressed in young elongating tissues.
==Annotated Information==

Brassinosteroids influence both plant height and leaf erectness in rice[1]. Arabidopsis thaliana BRASSINOSTEROID INSENSITIVE1 (bri1) mutants, which are deficient in the brassinosteroid receptor, have a severe dwarf phenotype[2]. The rice mutants d61-1 and d61-2, which are weak mutant alleles of OsBRI1, a homolog of A. thaliana BRI1, have both a semi-dwarf phenotype and more erect leaves[3]. In rice, an erect leaf phenotype has also been associated with loss-of-function, brassinosteroid-deficient mutants such as brassinosteroid-deficient dwarf1 (brd1), which is defective in OsDWARF (the homolog of tomato CYP85A1/DWARF[4]), and ebisu dwarf (d2), which is defective in CYP90D2/D2 (ref. [5]). However, the grain yield of d61, brd1 and d2 mutants is decreased because of morphological alterations in their reproductive development.

===Function===

New cultivars with very erect leaves, which increase light capture for photosynthesis and nitrogen storage for grain filling, may have increased grain yields.The erect leaf phenotype of a rice brassinosteroid-deficient mutant, osdwarf4-1, is associated with enhanced grain yields under conditions of dense planting, even without extra fertilizer. Molecular and biochemical studies reveal that two different cytochrome P450s, CYP90B2/OsDWARF4 and CYP724B1/D11, function redundantly in C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis. Therefore, despite the central role of brassinosteroids in plant growth and development, mutation of OsDWARF4 alone causes only limited defects in brassinosteroid biosynthesis and plant morphology[6].

===Expression===
The affected gene, OsDWARF4, was identified by characterizing the site of insertion of the Tos17 retrotransposon.

----Molecular characterization of OsDWARF4 and OsDWARF4L1/D11

OsDWARF4 is a homolog of A. thaliana DWARF4, which encodes a cytochrome P450, CYP90B1 (Fig. 1a). CYP90B1/DWARF4 catalyzes C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis[7]. Only one DWARF4 homolog was identified in the rice genome. OsDWARF4 is located on the short arm of chromosome 3 (35 cM). Its predicted open reading frame consists of eight exons, and encodes a protein of 502 amino acids (Fig. 1b,c). OsDWARF4 is most closely related to DWARF4 (66.3% amino-acid identity; Fig. 1a). The structure of OsDWARF4 is similar to that of DWARF4 throughout its length: five domains found in cytochrome P450s—proline rich, A (also referred to as dioxygen binding), B (steroid binding), C and heme binding—are highly conserved (Fig. 1c). In wild-type rice, OsDWARF4 transcripts are most abundant in leaf blades and roots, but are also found in all other plant parts tested (Fig. 1d). Transcript levels were decreased by exogenous brassinolide treatment, and increased in brassinosteroid-insensitive d61-3 and brassinosteroid-deficient brd1-1 (Fig. 1e). These results suggest that OsDWARF4 expression is feedback regulated, as is that of A. thaliana brassinosteroid-biosynthesis genes[8].
OsDWARF4L1 was identified as an OsDWARF4-like gene by in silico screening of rice DNA databases. OsDWARF4L1 is located on the long arm of chromosome 4 (70.9 cM). A recent study demonstrated that a rice semi-dwarf and short-grain mutant, d11, has a loss-of-function mutation in the OsDWARF4L1 gene[9]. OsDWARF4L1/D11 shows the highest homology with A. thaliana CYP724A1, and is closely related to rice OsDWARF4 and A. thaliana DWARF4 (47.4%, 37.7% and 41.7% amino-acid identities, respectively; Fig. 1a). In wild-type rice, OsDWARF4L1/D11 transcripts were most abundant in vegetative shoot apices, leaf sheaths and roots, and found at more moderate levels in other organs (Fig. 1d). They were decreased by exogenous brassinolide treatment, and increased in d61-3 and brd1-1 (Fig. 1e). These results indicate that OsDWARF4L1/D11 expression is regulated by a feedback mechanism, as is that of OsDWARF4.

----Phenotypes of rice mutants defective in brassinosteroid biosynthesis

In osdwarf4-1, Tos17 insertion into exon 6 causes a premature termination in domain C (arrowhead in Fig. 1b,c). Because a premature stop codon before the heme-binding domain caused loss-of-function of A. thaliana DWARF4 (ref. [7]), we consider osdwarf4-1 to be a null allele. Unexpectedly, osdwarf4-1 plants show only weak morphological phenotypes, such as a slightly dwarfed stature and more erect leaves (Fig. 2a,b). Both of these phenotypes were complemented by the introduction of an entire OsDWARF4 gene (data not shown). These results seem inconsistent with the fact that rice d61-3 and brd1-1, which are loss-of-function mutants of OsBRI1 and OsDWARF respectively, display severe dwarfing and have malformed leaves (for example, see brd1-1 in Fig. 2a), and suggest functional redundancy in brassinosteroid biosynthesis in rice.
Because null alleles of d11, such as d11-4, also show a weak phenotype[9] (Fig. 2a),OsDWARF4L1/D11 and OsDWARF4 may function redundantly. To confirm this possibility, researcher produced double mutants by crossing homozygous osdwarf4-1 and d11-4 plants. Among the F2 population, 34 of 576 plants showed severe dwarfing and malformed leaves with tortuous leaf blades, indistinguishable from the brassinosteroid-deficient brd1-1 phenotype (Fig. 2a,c,d). This ratio fits the theoretical 9:3:3:1 (wild-type:osdwarf4-1:d11-4:osdwarf4-1/d11-4) segregation predicted by mendelian inheritance. PCR and sequence analyses confirmed that these plants were osdwarf4-1/d11-4 double mutants (data not shown).
Endogenous levels of brassinosteroid intermediates in brd1 clearly indicate that OsDWARF catalyzes C-6 oxidation[4]. However, quantification in d2 could not determine the step(s) catalyzed by CYP90D2/D2, because CYP90D2/D2 functions redundantly with another gene, CYP90D3, and loss-of-function of CYP90D2/D2 have caused only limited defects in brassinosteroid biosynthesis[5]. Similarly, endogenous levels of brassinosteroid intermediates were obviously changed in the osdwarf4-1/d11-4 double mutant relative to osdwarf4-1 or d11 single mutants. Although brassinolide was not detected in either the wild-type or osdwarf4-1/d11-4, another bioactive brassinosteroid, castasterone (CS), was detected in the wild-type but not in osdwarf4-1/d11-4, confirming that osdwarf4-1/d11-4 is brassinosteroid-deficient (Fig. 2e). Levels of the other five intermediates downstream of C-22 hydroxylation—6-deoxocathasterone (6-DeoxoCT), 6-deoxoteasterone (6-DeoxoTE), 3-dehydro-6-deoxoteasterone (6-Deoxo3DT), 6-deoxotyphasterol (6-DeoxoTY) and 6-deoxocastasterone (6-DeoxoCS)—were also greatly reduced in osdwarf4-1/d11-4 (Fig. 2e). This indicates that both proteins are involved in C-22 hydroxylation. It is unclear why the upstream compound, campestanol, did not accumulate in osdwarf4-1/d11-4. Because A. thaliana CYP90B1/DWARF4 also catalyzes C-22 hydroxylation of other upstream compounds such as campesterol and (24R)-ergost-4-en-3-one[10], it is possible that campestanol is not the major substrate for CYP90B2/OsDWARF4 and CYP724B1/D11 in rice.

----Characterization of a field-grown osdwarf4-1 mutant

Because the harvest index (grain/(grain plus straw)) of rice is about 0.5, further increases in yield potential will have to involve increases in crop biomass driven by more net photosynthesis[11]. In rice, the contribution of lower leaves to photosynthesis is significant, even though their photosynthetic capacity is considerably less than that of upper leaves[12]. Erect leaves allow greater penetration of light to lower leaves, thereby optimizing canopy photosynthesis[13]. Therefore, the most likely explanation for the increase in above-ground biomass in osdwarf4-1 plots under dense planting conditions (more than 1.2 times that in wild-type under conventional planting condition) is that the shade of the upper leaves is minimized, and the lower leaves receive more light to drive higher rates of photosynthesis (Fig. 3 and Supplementary Table 2 online). No difference was observed in the heading date (flowering time) and the grain-filling period (time to maturity) between wild-type and osdwarf4-1 plants (data not shown). In the wild-type, the above-ground biomass was increased in response to higher fertilizer application under conventional planting (22.2 plants m-2), but was significantly decreased under dense planting with doubled fertilizer application owing to reduced photosynthesis caused by lodging (Fig. 3a). In contrast, the above-ground biomass of osdwarf4-1 was significantly (P < 0.01) increased by dense planting at each fertilizer rate. Consequently, the above-ground biomass in osdwarf4-1 plots under dense planting with standard and a 1.5-fold increase in fertilizer application was approx1.34 and 1.52 times that in wild-type plots under conventional cultivation conditions. The increased biomass depended mainly on accelerated formation of tillers, which have the ability to generate a panicle (panicle number, Fig. 3b). Both total and fertile grain numbers tended to increase in densely planted osdwarf4-1 plots (Fig. 3c), resulting in significantly increased grain yield even when the fertilizer application was not increased (for example, the osdwarf4-1 yield was 6.19 t ha-1 under dense planting with standard fertilizer application whereas the wild-type yield was 4.69 t ha-1 under conventional conditions; Fig. 3d). However, the increase in grain yield was not as much as that in above-ground biomass, because the harvest index of osdwarf4-1 (0.41) was slightly lower than that of the wild-type (0.44). A similar field test gave comparable results.

----Overexpression of OsDWARF4

Overexpression of DWARF in transgenic tomato increases plant height[14], and overexpression of DWARF4 in transgenic A. thaliana and tobacco increases vegetative growth and seed yield[15]. In rice, however, the role of brassinosteroid in regulating leaf angle has been known for over 40 years[16, 17]. Whereas brassinosteroid deficiency increased the erectness of leaves, the opposite phenotype (increased leaf angle) was observed in transgenic rice overexpressing OsDWARF4 (27.6° versus 10.0° in the wild-type, P < 0.01), and was inherited in their progeny (data not shown). Because inclined leaves are unfavorable for photosynthesis when plants are grown in a group, we have to carefully evaluate whether the overexpression of a brassinosteroid-biosynthesis gene in transgenic rice increases grain yield as is the case in transgenic plants overexpressing DWARF4.

===Conclusion===

More erect leaves improve not only light distribution but also the size and use of leaf nitrogen reservoirs for grain growth[13]. The erect leaf phenotype caused by brassinosteroid deficiency improved biomass production under field conditions, resulting in increased grain yield without the negative environmental effects associated with fertilizers. Recently identified rice homologs of A. thaliana BAS1 that encodes the brassinosteroid catabolic enzyme, CYP734A1 (ref. [18]). Overexpression of these rice homologs in transgenic rice successfully induced the brassinosteroid-deficient phenotypes (unpublished results). Although ideally, expression of these genes should be restricted to particular tissues and organs such as leaf laminae, it should be possible to generate erect leaf varieties by the introduction of a single transgene. In conclusion, engineering more erect leaves modulating brassinosteroid metabolism can be combined with dense planting to improve rice grain yields.

==Labs working on this gene==
Please input related labs here.

==References==
[1]Sakamoto T;Morinaka Y;Ohnishi T;Sunohara H;Fujioka S;Ueguchi Tanaka M;Mizutani M;Sakata K;Takatsuto S;Yoshida S;Tanaka H;Kitano H;Matsuoka M
Erect leaves caused by brassinosteroid deficiency increase biomass production and grain yield in rice
Nature Biotechnology, 2006, 24(1): 105-109.
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[2] Evans, L.T. Crop Evolution, Adaptation and Yield (Cambridge Univ. Press, Cambridge, 1993).
[3] Peng, J. et al. 'Green revolution' genes encode mutant gibberellin response modulators. Nature 400, 256–261 (1999).
[4] Sasaki, A. et al. Green revolution: a mutant gibberellin-synthesis gene in rice. Nature 416, 701–702 (2002).
[5] Sakamoto, T. et al. Genetic manipulation of gibberellin metabolism in transgenic rice. Nat. Biotechnol. 21, 909–913 (2003).
[6] Sakamoto, T. & Matsuoka, M. Generating high-yielding varieties by genetic manipulation of plant architecture. Curr. Opin. Biotech. 15, 144–147 (2004).
[7] Li, J. & Chory, J. A putative leucine-rich repeat receptor kinase involved in brassinosteroid signal transduction. Cell 90, 929–938 (1997).
[8] Yamamuro, C. et al. Loss of function of a rice brassinosteroid insensitive1 homolog prevents internode elongation and bending of the lamina joint. Plant Cell 12, 1591–1606 (2000).
[9] Hong, Z. et al. Loss-of-function of a rice brassinosteroid biosynthetic enzyme, C-6 oxidase, prevents the organized arrangement and polar elongation of cells in the leaves and stem. Plant J. 32, 495–508 (2002).
[10] Hong, Z. et al. A rice brassinosteroid-deficient mutant, ebisu dwarf (d2), is caused by a loss of function of a new member of cytochrome P450. Plant Cell 15, 2900–2910 (2003).
[11] Choe, S. et al. The DWF4 gene of Arabidopsis encodes a cytochrome P450 that mediates multiple 22alpha-hydroxylation steps in brassinosteroid biosynthesis. Plant Cell 10, 231–243 (1998).
[12] Bancos, S. et al. Regulation of transcript levels of the Arabidopsis cytochrome P450 genes involved in brassinosteroid biosynthesis. Plant Physiol. 130, 504–513 (2002).
[13] Tanabe, S. et al. A novel cytochrome P450 is implicated in brassinosteroid biosynthesis via the characterization of a rice dwarf mutant, dwarf11, with reduced seed length. Plant Cell 17, 776–790 (2005).
[14] Fujioka, S., Takatsuto, S. & Yoshida, S. An early C-22 oxidation branch in brassinosteroid biosynthetic pathway. Plant Physiol. 130, 930–939 (2002).
[15] Mann, C.C. Crop scientists seek a new revolution. Science 283, 310–314 (1999).
[16] Horton, P. Prospects for crop improvement through the genetic manipulation of photosynthesis: morphological and biochemical aspects of light capture. J. Exp. Bot. 51, 475–485 (2000).
[17] Bishop, G.J. et al. The tomato DWARF enzyme catalyses C-6 oxidation in brassinosteroid biosynthesis. Proc. Natl. Acad. Sci. USA 96, 1761–1766 (1999).
[18] Choe, S. et al. Overexpression of DWARF4 in the brassinosteroid biosynthetic pathway results in increased vegetative growth and seed yield in Arabidopsis. Plant J. 26, 573–582 (2001).
[19] Maeda, E. Rate of lamina inclination in excised rice leaves. Physiol. Plant. 18, 813–827 (1965).
[20] Wada, K. et al. Brassinolide and homobrassinolide promotion of lamina inclination of rice seedlings. Plant Cell Physiol. 22, 323–325 (1981).
[21] Neff, M.M. et al. BAS1: A gene regulating brassinosteroid levels and light responsiveness in Arabidopsis. Proc. Natl. Acad. Sci. USA 96, 15316–15323 (1999).
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[23] Saito, S. et al. Arabidopsis CYP707As encode (+)-abscisic acid 8'-hydroxylase, a key enzyme in the oxidative catabolism of abscisic acid. Plant Physiol. 134, 1439–1449 (2004).
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==Structured Information==
{{JaponicaGene|
GeneName = Os03g0227700|
Description = Cytochrome P450 family protein|
Version = NM_001055982.2 GI:297600576 GeneID:4332134|
Length = 6529 bp|
Definition = Oryza sativa Japonica Group Os03g0227700, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 3|Chromosome 3]]|
AP = Chromosome 3:6765109..6771637|
CDS = 6765109..6765374,6765543..6765867,6765971..6766123,6768937..6769188,6769337..6769429 ,6770222..6770300,6770398..6770504,6771392..6771637|
GCID = <gbrowseImage1>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</cdnaseq>|
AA = <aaseq>MAAMMASITSELLFFLPFILLALLTFYTTTVAKCHGGHWWRGGT TPAKRKRMNLPPGAAGWPLVGETFGYLRAHPATSVGRFMEQHIARYGKIYRSSLFGER TVVSADAGLNRYILQNEGRLFECSYPRSIGGILGKWSMLVLVGDPHREMRAISLNFLS SVRLRAVLLPEVERHTLLVLRAWPPSSTFSAQHQAKKFTFNLMAKNIMSMDPGEEETE RLRREYITFMKGVVSAPLNLPGTPYWKALKSRAAILGVIERKMEERVEKLSKEDASVE QDDLLGWALKQSNLSKEQILDLLLSLLFAGHETSSMALALAIFFLEGCPKAVQELREE HLGIARRQRLRGECKLSWEDYKEMVFTQCVINETLRLGNVVRFLHRKVIKDVHYKGYD IPSGWKILPVLAAVHLDSSLYEDPQRFNPWRWKSSGSSGGLAQSSSFMPYGGGTRLCA GSELAKLEMAVFLHHLVLNFRWELAEPDQAFVFPFVDFPKGLPIRVHRIAQDDEQE</aaseq>|
DNA = <dnaseqindica>1..266#435..759#863..1015#3829..4080#4229..4321#5114..5192#5290..5396#6284..6529#atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtcggtggtttcattactcggtacacgcacacagctatataagcacgcgcactgctcgcctcgccgtgaagcaagcttagctagtagctaggatatcgatcgatcggtaataataggctgacggcgagctggggggtggcgtgcggttgcgtgcgtgcgtgcatgcaggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaaggtacaacacacatcgtcctccattaagttgagcatgcatatatcaatcagctcgctctagcgatcaatatcaatgaaacgcgcgtttgcgtgcgtgcgtgcagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaaggtacgtactcgtagctactagctactccctccgtttcgaaatgtttgacgccgttgactttttatcacatatttgaccgttcgttttatttaaaaaaattaagtaattattaattattttcctatcatttgattcattgttaaatatatttttatgtaggcatataattttacatatctcacaaaagtttttaaataagacgaacggtcaaacatttgctaaaaagtcaacggtgtcaaatatttcgaaacggaggaagtagctagctaccaccccacccgtgcggattgatcgatcgatcgatgtccctctctgcctacacacaccctgcgccccggccgtacgcgcgcaaagtcgtgcgcgcatgagttttactccttccgtttcaaaaaaaaaacgacctagacaagtaatatgatgtaacctagaacaacgaatctatacaggagtatatccaaattcattgtcctagattacatcatatctttgtctatgttgccttttttaggaggagagagtactttacatatgagttatgaccggccggccaggttcctttcttcccctattccgtatgcatactataagcagctgcatataggcatatactactatacccagcttagctaaagcaaacaacttgcaacgacgatcatgtgcaggtccctgttgacagaggttagagatgtaaaccatttttgttgttttcattatctaaagagaaaacaagttactacgacgatcgagatgtacgcaaggggatcatatttatattggagaggagagagagatgatttgcgacttgtgagtgataagtgaatatatatactatatatttgtgtaggagtgacgatgacttccctgtcaatgttttgtttggtatgcacacggggggaaaagagtttagagaaaggaattggctagagagggccaggtttagtttctttaatttttgtttggtggttttgtgtgtagctagtccgcatgcatgatatatcatatatatgcatggcccatcattagatatatatacatgacggcagtgcacttattatatatatgtaggggcatatacaaggtgtatgtgtttagaattgtggatggtccaggaattgttgcagctagttctatactagcatagtgaattagtgatccatatcgagatcaggtccaatcggtcagctcgcacagcgggtcttttagtacgatttgttcggacaaattatcttgcctttgagataatcaacataggattgaaactgcatactagcaatagaccatagctatttggcaatatagaaagctaaaaaatgttgattgaaaggtgcgtgcatgcatgctagcagtatgtcttttaggccaggtccagtttttttttttttctcaaaggcagttgcttaaggtgaaattctagcagaggtcatatgaatatatttattatcttccaaacatgagccaaaacctatatatagttaactcaaaaaaccatgcattggtacttcttctgagtatgcatcctggggactgatcgatcatgcagattcttttctttctttttccctggggacgtactgatcatcactcatgtgagcataaaatgttgcctactttttactgcatgtgaggtcacactcaaatgatctttattgattgcctcagcatgcatgctgctagctttcacgatctctctccgatacacacaccctgtgctgtgctcaacagcatgcaacctggcaggtggaatttttaatctgttggaaagtaaatgaccaaccttcataatttggcatgattatatgctggggaatctatcatttatcatgattcaaccgctgaccaaactctttgccacacacccgaaccacattaatttcgttaagcggatgattagccagcaaccttcacagtttggttttaattagttaatgaaagggtgcttgtagttttattagctgaagagacactgcagtactcctactgagcgttctccttaattgggaaatagatctctgcagtactatcctttaactttaagtgaataacatgtagctaggtctcacacttatcaggcccacacatcagtgatccaagtcaaatggcaatacaccataggagattcttccactaattggctaacaaaaataaactgtgatcactgatcacagctgctaacactataacacattaatttgcatgtctcatgaaatgccagatgagttggattaattatctatcttatgttcagtgccgaggaatatacaaggaaacatcattggtatagttgtggagttctgagtgcattatctcacccagtacggtcaaattaatcctagtctccgtaaatatggttgtgtgcactcttagttaattagtgtagacccggaaattatactttgtcgagaatgttgttatccgtgtgttaagtttcaagtaccctgcaggcctataactaaacactgccaacatggatgtttaagaatgaattatacatacagcgcacttggaaaagaaaaggggtagctacccagctgaagaacacctctctctcatttgaaggtcgacacaatatgtgtcttttctatatcgaaaggaaacgacattttttctcttcaaaacagaaagctatgacaagagacataatcctagactagtaggcggttaggcaacaccatatctgaatttgataggtgtcccacatattttttttcttttcctgataagaaggaacttttcaccgtgagcttatttttttctgttctatgtcgacattagttccatgatgtttactgtgtatttttactgttcccctgcagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggtacagagaattacagattcgaaattaacatctactgtcactagctcagtgctacgtttaactgtagagcaacaattgcagaaagcgaaggcatcagaaatgacatgggaatcgattagttacactcacactgacgcgtgacatgcaggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtaaggccccccctactttgttccttccgatccctcctccctaaacgctactaatataaatcggtgcaaaggctttgtgctagcgtagctcttgctgttttgtttggtcacttgcgtagagagatagcaacttcccgaaatgtcatgtacgaatggagactgaaagctctgcgaaataattgccgggccacgtactacagtacttacgctccctctcttcagtttggatcgatcgagctaagaaggcgcacgcatgtgtgcattgatcgatagagtcgatctgttctgccttgtaagctagccagctgaggttctttcgccttggaagtttctacagggtgaggcccatcgtcttgcaacgatccaaatgccatgacaagtacctgttgttgtaggcttccagtagcacggcctgcccccaagaaggtatctgaatcttctgcattgctgttcagacggcagagcagaacatgccgaaataccattccagtcagttgcagtatatgtgtgtggtggtatcagataaaggagcagagtaatttgaggccttagaaattcagatgagttgctcgacaattgtcgcc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Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001055982.2 RefSeq:Os03g0227700]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 3]]
[[Category:Chromosome 3]]

File:搜狗截图14年05月24日2044 1.png

2014-05-24T13:10:14Z

LQ:

Os03g0227700

2014-05-24T13:07:22Z

LQ: /* References */

Please input one-sentence summary here.
''OsDWAR''F4 is expressed in young elongating tissues.
==Annotated Information==

Brassinosteroids influence both plant height and leaf erectness in rice[6]. Arabidopsis thaliana BRASSINOSTEROID INSENSITIVE1 (bri1) mutants, which are deficient in the brassinosteroid receptor, have a severe dwarf phenotype[7]. The rice mutants d61-1 and d61-2, which are weak mutant alleles of OsBRI1, a homolog of A. thaliana BRI1, have both a semi-dwarf phenotype and more erect leaves[8]. In rice, an erect leaf phenotype has also been associated with loss-of-function, brassinosteroid-deficient mutants such as brassinosteroid-deficient dwarf1 (brd1), which is defective in OsDWARF (the homolog of tomato CYP85A1/DWARF[9]), and ebisu dwarf (d2), which is defective in CYP90D2/D2 (ref. [10]). However, the grain yield of d61, brd1 and d2 mutants is decreased because of morphological alterations in their reproductive development.

===Function===

New cultivars with very erect leaves, which increase light capture for photosynthesis and nitrogen storage for grain filling, may have increased grain yields.The erect leaf phenotype of a rice brassinosteroid-deficient mutant, osdwarf4-1, is associated with enhanced grain yields under conditions of dense planting, even without extra fertilizer. Molecular and biochemical studies reveal that two different cytochrome P450s, CYP90B2/OsDWARF4 and CYP724B1/D11, function redundantly in C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis. Therefore, despite the central role of brassinosteroids in plant growth and development, mutation of OsDWARF4 alone causes only limited defects in brassinosteroid biosynthesis and plant morphology[1].

===Expression===
The affected gene, OsDWARF4, was identified by characterizing the site of insertion of the Tos17 retrotransposon.

----Molecular characterization of OsDWARF4 and OsDWARF4L1/D11

OsDWARF4 is a homolog of A. thaliana DWARF4, which encodes a cytochrome P450, CYP90B1 (Fig. 1a). CYP90B1/DWARF4 catalyzes C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis[11]. Only one DWARF4 homolog was identified in the rice genome. OsDWARF4 is located on the short arm of chromosome 3 (35 cM). Its predicted open reading frame consists of eight exons, and encodes a protein of 502 amino acids (Fig. 1b,c). OsDWARF4 is most closely related to DWARF4 (66.3% amino-acid identity; Fig. 1a). The structure of OsDWARF4 is similar to that of DWARF4 throughout its length: five domains found in cytochrome P450s—proline rich, A (also referred to as dioxygen binding), B (steroid binding), C and heme binding—are highly conserved (Fig. 1c). In wild-type rice, OsDWARF4 transcripts are most abundant in leaf blades and roots, but are also found in all other plant parts tested (Fig. 1d). Transcript levels were decreased by exogenous brassinolide treatment, and increased in brassinosteroid-insensitive d61-3 and brassinosteroid-deficient brd1-1 (Fig. 1e). These results suggest that OsDWARF4 expression is feedback regulated, as is that of A. thaliana brassinosteroid-biosynthesis genes[12].
OsDWARF4L1 was identified as an OsDWARF4-like gene by in silico screening of rice DNA databases. OsDWARF4L1 is located on the long arm of chromosome 4 (70.9 cM). A recent study demonstrated that a rice semi-dwarf and short-grain mutant, d11, has a loss-of-function mutation in the OsDWARF4L1 gene13. OsDWARF4L1/D11 shows the highest homology with A. thaliana CYP724A1, and is closely related to rice OsDWARF4 and A. thaliana DWARF4 (47.4%, 37.7% and 41.7% amino-acid identities, respectively; Fig. 1a). In wild-type rice, OsDWARF4L1/D11 transcripts were most abundant in vegetative shoot apices, leaf sheaths and roots, and found at more moderate levels in other organs (Fig. 1d). They were decreased by exogenous brassinolide treatment, and increased in d61-3 and brd1-1 (Fig. 1e). These results indicate that OsDWARF4L1/D11 expression is regulated by a feedback mechanism, as is that of OsDWARF4.

----Phenotypes of rice mutants defective in brassinosteroid biosynthesis

In osdwarf4-1, Tos17 insertion into exon 6 causes a premature termination in domain C (arrowhead in Fig. 1b,c). Because a premature stop codon before the heme-binding domain caused loss-of-function of A. thaliana DWARF4 (ref. 11), we consider osdwarf4-1 to be a null allele. Unexpectedly, osdwarf4-1 plants show only weak morphological phenotypes, such as a slightly dwarfed stature and more erect leaves (Fig. 2a,b). Both of these phenotypes were complemented by the introduction of an entire OsDWARF4 gene (data not shown). These results seem inconsistent with the fact that rice d61-3 and brd1-1, which are loss-of-function mutants of OsBRI1 and OsDWARF respectively, display severe dwarfing and have malformed leaves (for example, see brd1-1 in Fig. 2a), and suggest functional redundancy in brassinosteroid biosynthesis in rice.
Because null alleles of d11, such as d11-4, also show a weak phenotype13 (Fig. 2a), we suspected that OsDWARF4L1/D11 and OsDWARF4 may function redundantly. To confirm this possibility, we produced double mutants by crossing homozygous osdwarf4-1 and d11-4 plants. Among the F2 population, 34 of 576 plants showed severe dwarfing and malformed leaves with tortuous leaf blades, indistinguishable from the brassinosteroid-deficient brd1-1 phenotype (Fig. 2a,c,d). This ratio fits the theoretical 9:3:3:1 (wild-type:osdwarf4-1:d11-4:osdwarf4-1/d11-4) segregation predicted by mendelian inheritance. PCR and sequence analyses confirmed that these plants were osdwarf4-1/d11-4 double mutants (data not shown).
Endogenous levels of brassinosteroid intermediates in brd1 clearly indicate that OsDWARF catalyzes C-6 oxidation9. However, quantification in d2 could not determine the step(s) catalyzed by CYP90D2/D2, because CYP90D2/D2 functions redundantly with another gene, CYP90D3, and loss-of-function of CYP90D2/D2 have caused only limited defects in brassinosteroid biosynthesis10. Similarly, endogenous levels of brassinosteroid intermediates were obviously changed in the osdwarf4-1/d11-4 double mutant relative to osdwarf4-1 or d11 single mutants. Although brassinolide was not detected in either the wild-type or osdwarf4-1/d11-4, another bioactive brassinosteroid, castasterone (CS), was detected in the wild-type but not in osdwarf4-1/d11-4, confirming that osdwarf4-1/d11-4 is brassinosteroid-deficient (Fig. 2e). Levels of the other five intermediates downstream of C-22 hydroxylation—6-deoxocathasterone (6-DeoxoCT), 6-deoxoteasterone (6-DeoxoTE), 3-dehydro-6-deoxoteasterone (6-Deoxo3DT), 6-deoxotyphasterol (6-DeoxoTY) and 6-deoxocastasterone (6-DeoxoCS)—were also greatly reduced in osdwarf4-1/d11-4 (Fig. 2e). This indicates that both proteins are involved in C-22 hydroxylation. It is unclear why the upstream compound, campestanol, did not accumulate in osdwarf4-1/d11-4. Because A. thaliana CYP90B1/DWARF4 also catalyzes C-22 hydroxylation of other upstream compounds such as campesterol and (24R)-ergost-4-en-3-one14, it is possible that campestanol is not the major substrate for CYP90B2/OsDWARF4 and CYP724B1/D11 in rice.

----Characterization of a field-grown osdwarf4-1 mutant

Because the harvest index (grain/(grain plus straw)) of rice is about 0.5, further increases in yield potential will have to involve increases in crop biomass driven by more net photosynthesis15. In rice, the contribution of lower leaves to photosynthesis is significant, even though their photosynthetic capacity is considerably less than that of upper leaves16. Erect leaves allow greater penetration of light to lower leaves, thereby optimizing canopy photosynthesis1. Therefore, the most likely explanation for the increase in above-ground biomass in osdwarf4-1 plots under dense planting conditions (more than 1.2 times that in wild-type under conventional planting condition) is that the shade of the upper leaves is minimized, and the lower leaves receive more light to drive higher rates of photosynthesis (Fig. 3 and Supplementary Table 2 online). No difference was observed in the heading date (flowering time) and the grain-filling period (time to maturity) between wild-type and osdwarf4-1 plants (data not shown). In the wild-type, the above-ground biomass was increased in response to higher fertilizer application under conventional planting (22.2 plants m-2), but was significantly decreased under dense planting with doubled fertilizer application owing to reduced photosynthesis caused by lodging (Fig. 3a). In contrast, the above-ground biomass of osdwarf4-1 was significantly (P < 0.01) increased by dense planting at each fertilizer rate. Consequently, the above-ground biomass in osdwarf4-1 plots under dense planting with standard and a 1.5-fold increase in fertilizer application was approx1.34 and 1.52 times that in wild-type plots under conventional cultivation conditions. The increased biomass depended mainly on accelerated formation of tillers, which have the ability to generate a panicle (panicle number, Fig. 3b). Both total and fertile grain numbers tended to increase in densely planted osdwarf4-1 plots (Fig. 3c), resulting in significantly increased grain yield even when the fertilizer application was not increased (for example, the osdwarf4-1 yield was 6.19 t ha-1 under dense planting with standard fertilizer application whereas the wild-type yield was 4.69 t ha-1 under conventional conditions; Fig. 3d). However, the increase in grain yield was not as much as that in above-ground biomass, because the harvest index of osdwarf4-1 (0.41) was slightly lower than that of the wild-type (0.44). A similar field test gave comparable results.

----Overexpression of OsDWARF4

Overexpression of DWARF in transgenic tomato increases plant height17, and overexpression of DWARF4 in transgenic A. thaliana and tobacco increases vegetative growth and seed yield18. In rice, however, the role of brassinosteroid in regulating leaf angle has been known for over 40 years19, 20. Whereas brassinosteroid deficiency increased the erectness of leaves, the opposite phenotype (increased leaf angle) was observed in transgenic rice overexpressing OsDWARF4 (27.6° versus 10.0° in the wild-type, P < 0.01), and was inherited in their progeny (data not shown). Because inclined leaves are unfavorable for photosynthesis when plants are grown in a group, we have to carefully evaluate whether the overexpression of a brassinosteroid-biosynthesis gene in transgenic rice increases grain yield as is the case in transgenic plants overexpressing DWARF4.

===Conclusion===

A previous study concluded that more erect leaves improve not only light distribution but also the size and use of leaf nitrogen reservoirs for grain growth1. Our results clearly demonstrate that the erect leaf phenotype caused by brassinosteroid deficiency improved biomass production under field conditions, resulting in increased grain yield without the negative environmental effects associated with fertilizers. We have recently identified rice homologs of A. thaliana BAS1 that encodes the brassinosteroid catabolic enzyme, CYP734A1 (ref. 21). Overexpression of these rice homologs in transgenic rice successfully induced the brassinosteroid-deficient phenotypes (unpublished results). Although ideally, expression of these genes should be restricted to particular tissues and organs such as leaf laminae, it should be possible to generate erect leaf varieties by the introduction of a single transgene. In conclusion, engineering more erect leaves modulating brassinosteroid metabolism can be combined with dense planting to improve rice grain yields.

==Labs working on this gene==
Please input related labs here.

==References==
[1]Sakamoto T;Morinaka Y;Ohnishi T;Sunohara H;Fujioka S;Ueguchi Tanaka M;Mizutani M;Sakata K;Takatsuto S;Yoshida S;Tanaka H;Kitano H;Matsuoka M
Erect leaves caused by brassinosteroid deficiency increase biomass production and grain yield in rice
Nature Biotechnology, 2006, 24(1): 105-109.
[1] Sinclair, T.R. & Sheehy, J.E. Erect leaves and photosynthesis in rice. Science 283, 1455 (1999).
[2] Evans, L.T. Crop Evolution, Adaptation and Yield (Cambridge Univ. Press, Cambridge, 1993).
[3] Peng, J. et al. 'Green revolution' genes encode mutant gibberellin response modulators. Nature 400, 256–261 (1999).
[4] Sasaki, A. et al. Green revolution: a mutant gibberellin-synthesis gene in rice. Nature 416, 701–702 (2002).
[5] Sakamoto, T. et al. Genetic manipulation of gibberellin metabolism in transgenic rice. Nat. Biotechnol. 21, 909–913 (2003).
[6] Sakamoto, T. & Matsuoka, M. Generating high-yielding varieties by genetic manipulation of plant architecture. Curr. Opin. Biotech. 15, 144–147 (2004).
[7] Li, J. & Chory, J. A putative leucine-rich repeat receptor kinase involved in brassinosteroid signal transduction. Cell 90, 929–938 (1997).
[8] Yamamuro, C. et al. Loss of function of a rice brassinosteroid insensitive1 homolog prevents internode elongation and bending of the lamina joint. Plant Cell 12, 1591–1606 (2000).
[9] Hong, Z. et al. Loss-of-function of a rice brassinosteroid biosynthetic enzyme, C-6 oxidase, prevents the organized arrangement and polar elongation of cells in the leaves and stem. Plant J. 32, 495–508 (2002).
[10] Hong, Z. et al. A rice brassinosteroid-deficient mutant, ebisu dwarf (d2), is caused by a loss of function of a new member of cytochrome P450. Plant Cell 15, 2900–2910 (2003).
[11] Choe, S. et al. The DWF4 gene of Arabidopsis encodes a cytochrome P450 that mediates multiple 22alpha-hydroxylation steps in brassinosteroid biosynthesis. Plant Cell 10, 231–243 (1998).
[12] Bancos, S. et al. Regulation of transcript levels of the Arabidopsis cytochrome P450 genes involved in brassinosteroid biosynthesis. Plant Physiol. 130, 504–513 (2002).
[13] Tanabe, S. et al. A novel cytochrome P450 is implicated in brassinosteroid biosynthesis via the characterization of a rice dwarf mutant, dwarf11, with reduced seed length. Plant Cell 17, 776–790 (2005).
[14] Fujioka, S., Takatsuto, S. & Yoshida, S. An early C-22 oxidation branch in brassinosteroid biosynthetic pathway. Plant Physiol. 130, 930–939 (2002).
[15] Mann, C.C. Crop scientists seek a new revolution. Science 283, 310–314 (1999).
[16] Horton, P. Prospects for crop improvement through the genetic manipulation of photosynthesis: morphological and biochemical aspects of light capture. J. Exp. Bot. 51, 475–485 (2000).
[17] Bishop, G.J. et al. The tomato DWARF enzyme catalyses C-6 oxidation in brassinosteroid biosynthesis. Proc. Natl. Acad. Sci. USA 96, 1761–1766 (1999).
[18] Choe, S. et al. Overexpression of DWARF4 in the brassinosteroid biosynthetic pathway results in increased vegetative growth and seed yield in Arabidopsis. Plant J. 26, 573–582 (2001).
[19] Maeda, E. Rate of lamina inclination in excised rice leaves. Physiol. Plant. 18, 813–827 (1965).
[20] Wada, K. et al. Brassinolide and homobrassinolide promotion of lamina inclination of rice seedlings. Plant Cell Physiol. 22, 323–325 (1981).
[21] Neff, M.M. et al. BAS1: A gene regulating brassinosteroid levels and light responsiveness in Arabidopsis. Proc. Natl. Acad. Sci. USA 96, 15316–15323 (1999).
[22] Sakamoto, T. et al. An overview of gibberellin metabolism enzyme genes and their related mutants in rice. Plant Physiol. 134, 1642–1653 (2004).
[23] Saito, S. et al. Arabidopsis CYP707As encode (+)-abscisic acid 8'-hydroxylase, a key enzyme in the oxidative catabolism of abscisic acid. Plant Physiol. 134, 1439–1449 (2004).
[24] Fujita, S. et al. Arabidopsis CYP90B1 catalyzes the early C-22 hydroxylation of C27, C28, and C29 sterols. Plant J. 45, in press.
[25] Hiei, Y., Ohta, S., Komari, T. & Kumashiro, T. Efficient transformation of rice (Oryza sativa L.) mediated by Agrobacterium and sequence analysis of boundaries of the T-DNA. Plant J. 6, 271–282 (1994).

==Structured Information==
{{JaponicaGene|
GeneName = Os03g0227700|
Description = Cytochrome P450 family protein|
Version = NM_001055982.2 GI:297600576 GeneID:4332134|
Length = 6529 bp|
Definition = Oryza sativa Japonica Group Os03g0227700, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 3|Chromosome 3]]|
AP = Chromosome 3:6765109..6771637|
CDS = 6765109..6765374,6765543..6765867,6765971..6766123,6768937..6769188,6769337..6769429 ,6770222..6770300,6770398..6770504,6771392..6771637|
GCID = <gbrowseImage1>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</cdnaseq>|
AA = <aaseq>MAAMMASITSELLFFLPFILLALLTFYTTTVAKCHGGHWWRGGT TPAKRKRMNLPPGAAGWPLVGETFGYLRAHPATSVGRFMEQHIARYGKIYRSSLFGER TVVSADAGLNRYILQNEGRLFECSYPRSIGGILGKWSMLVLVGDPHREMRAISLNFLS SVRLRAVLLPEVERHTLLVLRAWPPSSTFSAQHQAKKFTFNLMAKNIMSMDPGEEETE RLRREYITFMKGVVSAPLNLPGTPYWKALKSRAAILGVIERKMEERVEKLSKEDASVE QDDLLGWALKQSNLSKEQILDLLLSLLFAGHETSSMALALAIFFLEGCPKAVQELREE HLGIARRQRLRGECKLSWEDYKEMVFTQCVINETLRLGNVVRFLHRKVIKDVHYKGYD IPSGWKILPVLAAVHLDSSLYEDPQRFNPWRWKSSGSSGGLAQSSSFMPYGGGTRLCA GSELAKLEMAVFLHHLVLNFRWELAEPDQAFVFPFVDFPKGLPIRVHRIAQDDEQE</aaseq>|
DNA = 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Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001055982.2 RefSeq:Os03g0227700]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 3]]
[[Category:Chromosome 3]]

Os03g0227700

2014-05-24T13:01:24Z

LQ: /* Annotated Information */

Please input one-sentence summary here.
''OsDWAR''F4 is expressed in young elongating tissues.
==Annotated Information==

Brassinosteroids influence both plant height and leaf erectness in rice[6]. Arabidopsis thaliana BRASSINOSTEROID INSENSITIVE1 (bri1) mutants, which are deficient in the brassinosteroid receptor, have a severe dwarf phenotype[7]. The rice mutants d61-1 and d61-2, which are weak mutant alleles of OsBRI1, a homolog of A. thaliana BRI1, have both a semi-dwarf phenotype and more erect leaves[8]. In rice, an erect leaf phenotype has also been associated with loss-of-function, brassinosteroid-deficient mutants such as brassinosteroid-deficient dwarf1 (brd1), which is defective in OsDWARF (the homolog of tomato CYP85A1/DWARF[9]), and ebisu dwarf (d2), which is defective in CYP90D2/D2 (ref. [10]). However, the grain yield of d61, brd1 and d2 mutants is decreased because of morphological alterations in their reproductive development.

===Function===

New cultivars with very erect leaves, which increase light capture for photosynthesis and nitrogen storage for grain filling, may have increased grain yields.The erect leaf phenotype of a rice brassinosteroid-deficient mutant, osdwarf4-1, is associated with enhanced grain yields under conditions of dense planting, even without extra fertilizer. Molecular and biochemical studies reveal that two different cytochrome P450s, CYP90B2/OsDWARF4 and CYP724B1/D11, function redundantly in C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis. Therefore, despite the central role of brassinosteroids in plant growth and development, mutation of OsDWARF4 alone causes only limited defects in brassinosteroid biosynthesis and plant morphology[1].

===Expression===
The affected gene, OsDWARF4, was identified by characterizing the site of insertion of the Tos17 retrotransposon.

----Molecular characterization of OsDWARF4 and OsDWARF4L1/D11

OsDWARF4 is a homolog of A. thaliana DWARF4, which encodes a cytochrome P450, CYP90B1 (Fig. 1a). CYP90B1/DWARF4 catalyzes C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis[11]. Only one DWARF4 homolog was identified in the rice genome. OsDWARF4 is located on the short arm of chromosome 3 (35 cM). Its predicted open reading frame consists of eight exons, and encodes a protein of 502 amino acids (Fig. 1b,c). OsDWARF4 is most closely related to DWARF4 (66.3% amino-acid identity; Fig. 1a). The structure of OsDWARF4 is similar to that of DWARF4 throughout its length: five domains found in cytochrome P450s—proline rich, A (also referred to as dioxygen binding), B (steroid binding), C and heme binding—are highly conserved (Fig. 1c). In wild-type rice, OsDWARF4 transcripts are most abundant in leaf blades and roots, but are also found in all other plant parts tested (Fig. 1d). Transcript levels were decreased by exogenous brassinolide treatment, and increased in brassinosteroid-insensitive d61-3 and brassinosteroid-deficient brd1-1 (Fig. 1e). These results suggest that OsDWARF4 expression is feedback regulated, as is that of A. thaliana brassinosteroid-biosynthesis genes[12].
OsDWARF4L1 was identified as an OsDWARF4-like gene by in silico screening of rice DNA databases. OsDWARF4L1 is located on the long arm of chromosome 4 (70.9 cM). A recent study demonstrated that a rice semi-dwarf and short-grain mutant, d11, has a loss-of-function mutation in the OsDWARF4L1 gene13. OsDWARF4L1/D11 shows the highest homology with A. thaliana CYP724A1, and is closely related to rice OsDWARF4 and A. thaliana DWARF4 (47.4%, 37.7% and 41.7% amino-acid identities, respectively; Fig. 1a). In wild-type rice, OsDWARF4L1/D11 transcripts were most abundant in vegetative shoot apices, leaf sheaths and roots, and found at more moderate levels in other organs (Fig. 1d). They were decreased by exogenous brassinolide treatment, and increased in d61-3 and brd1-1 (Fig. 1e). These results indicate that OsDWARF4L1/D11 expression is regulated by a feedback mechanism, as is that of OsDWARF4.

----Phenotypes of rice mutants defective in brassinosteroid biosynthesis

In osdwarf4-1, Tos17 insertion into exon 6 causes a premature termination in domain C (arrowhead in Fig. 1b,c). Because a premature stop codon before the heme-binding domain caused loss-of-function of A. thaliana DWARF4 (ref. 11), we consider osdwarf4-1 to be a null allele. Unexpectedly, osdwarf4-1 plants show only weak morphological phenotypes, such as a slightly dwarfed stature and more erect leaves (Fig. 2a,b). Both of these phenotypes were complemented by the introduction of an entire OsDWARF4 gene (data not shown). These results seem inconsistent with the fact that rice d61-3 and brd1-1, which are loss-of-function mutants of OsBRI1 and OsDWARF respectively, display severe dwarfing and have malformed leaves (for example, see brd1-1 in Fig. 2a), and suggest functional redundancy in brassinosteroid biosynthesis in rice.
Because null alleles of d11, such as d11-4, also show a weak phenotype13 (Fig. 2a), we suspected that OsDWARF4L1/D11 and OsDWARF4 may function redundantly. To confirm this possibility, we produced double mutants by crossing homozygous osdwarf4-1 and d11-4 plants. Among the F2 population, 34 of 576 plants showed severe dwarfing and malformed leaves with tortuous leaf blades, indistinguishable from the brassinosteroid-deficient brd1-1 phenotype (Fig. 2a,c,d). This ratio fits the theoretical 9:3:3:1 (wild-type:osdwarf4-1:d11-4:osdwarf4-1/d11-4) segregation predicted by mendelian inheritance. PCR and sequence analyses confirmed that these plants were osdwarf4-1/d11-4 double mutants (data not shown).
Endogenous levels of brassinosteroid intermediates in brd1 clearly indicate that OsDWARF catalyzes C-6 oxidation9. However, quantification in d2 could not determine the step(s) catalyzed by CYP90D2/D2, because CYP90D2/D2 functions redundantly with another gene, CYP90D3, and loss-of-function of CYP90D2/D2 have caused only limited defects in brassinosteroid biosynthesis10. Similarly, endogenous levels of brassinosteroid intermediates were obviously changed in the osdwarf4-1/d11-4 double mutant relative to osdwarf4-1 or d11 single mutants. Although brassinolide was not detected in either the wild-type or osdwarf4-1/d11-4, another bioactive brassinosteroid, castasterone (CS), was detected in the wild-type but not in osdwarf4-1/d11-4, confirming that osdwarf4-1/d11-4 is brassinosteroid-deficient (Fig. 2e). Levels of the other five intermediates downstream of C-22 hydroxylation—6-deoxocathasterone (6-DeoxoCT), 6-deoxoteasterone (6-DeoxoTE), 3-dehydro-6-deoxoteasterone (6-Deoxo3DT), 6-deoxotyphasterol (6-DeoxoTY) and 6-deoxocastasterone (6-DeoxoCS)—were also greatly reduced in osdwarf4-1/d11-4 (Fig. 2e). This indicates that both proteins are involved in C-22 hydroxylation. It is unclear why the upstream compound, campestanol, did not accumulate in osdwarf4-1/d11-4. Because A. thaliana CYP90B1/DWARF4 also catalyzes C-22 hydroxylation of other upstream compounds such as campesterol and (24R)-ergost-4-en-3-one14, it is possible that campestanol is not the major substrate for CYP90B2/OsDWARF4 and CYP724B1/D11 in rice.

----Characterization of a field-grown osdwarf4-1 mutant

Because the harvest index (grain/(grain plus straw)) of rice is about 0.5, further increases in yield potential will have to involve increases in crop biomass driven by more net photosynthesis15. In rice, the contribution of lower leaves to photosynthesis is significant, even though their photosynthetic capacity is considerably less than that of upper leaves16. Erect leaves allow greater penetration of light to lower leaves, thereby optimizing canopy photosynthesis1. Therefore, the most likely explanation for the increase in above-ground biomass in osdwarf4-1 plots under dense planting conditions (more than 1.2 times that in wild-type under conventional planting condition) is that the shade of the upper leaves is minimized, and the lower leaves receive more light to drive higher rates of photosynthesis (Fig. 3 and Supplementary Table 2 online). No difference was observed in the heading date (flowering time) and the grain-filling period (time to maturity) between wild-type and osdwarf4-1 plants (data not shown). In the wild-type, the above-ground biomass was increased in response to higher fertilizer application under conventional planting (22.2 plants m-2), but was significantly decreased under dense planting with doubled fertilizer application owing to reduced photosynthesis caused by lodging (Fig. 3a). In contrast, the above-ground biomass of osdwarf4-1 was significantly (P < 0.01) increased by dense planting at each fertilizer rate. Consequently, the above-ground biomass in osdwarf4-1 plots under dense planting with standard and a 1.5-fold increase in fertilizer application was approx1.34 and 1.52 times that in wild-type plots under conventional cultivation conditions. The increased biomass depended mainly on accelerated formation of tillers, which have the ability to generate a panicle (panicle number, Fig. 3b). Both total and fertile grain numbers tended to increase in densely planted osdwarf4-1 plots (Fig. 3c), resulting in significantly increased grain yield even when the fertilizer application was not increased (for example, the osdwarf4-1 yield was 6.19 t ha-1 under dense planting with standard fertilizer application whereas the wild-type yield was 4.69 t ha-1 under conventional conditions; Fig. 3d). However, the increase in grain yield was not as much as that in above-ground biomass, because the harvest index of osdwarf4-1 (0.41) was slightly lower than that of the wild-type (0.44). A similar field test gave comparable results.

----Overexpression of OsDWARF4

Overexpression of DWARF in transgenic tomato increases plant height17, and overexpression of DWARF4 in transgenic A. thaliana and tobacco increases vegetative growth and seed yield18. In rice, however, the role of brassinosteroid in regulating leaf angle has been known for over 40 years19, 20. Whereas brassinosteroid deficiency increased the erectness of leaves, the opposite phenotype (increased leaf angle) was observed in transgenic rice overexpressing OsDWARF4 (27.6° versus 10.0° in the wild-type, P < 0.01), and was inherited in their progeny (data not shown). Because inclined leaves are unfavorable for photosynthesis when plants are grown in a group, we have to carefully evaluate whether the overexpression of a brassinosteroid-biosynthesis gene in transgenic rice increases grain yield as is the case in transgenic plants overexpressing DWARF4.

===Conclusion===

A previous study concluded that more erect leaves improve not only light distribution but also the size and use of leaf nitrogen reservoirs for grain growth1. Our results clearly demonstrate that the erect leaf phenotype caused by brassinosteroid deficiency improved biomass production under field conditions, resulting in increased grain yield without the negative environmental effects associated with fertilizers. We have recently identified rice homologs of A. thaliana BAS1 that encodes the brassinosteroid catabolic enzyme, CYP734A1 (ref. 21). Overexpression of these rice homologs in transgenic rice successfully induced the brassinosteroid-deficient phenotypes (unpublished results). Although ideally, expression of these genes should be restricted to particular tissues and organs such as leaf laminae, it should be possible to generate erect leaf varieties by the introduction of a single transgene. In conclusion, engineering more erect leaves modulating brassinosteroid metabolism can be combined with dense planting to improve rice grain yields.

==Labs working on this gene==
Please input related labs here.

==References==
1. Sakamoto T;Morinaka Y;Ohnishi T;Sunohara H;Fujioka S;Ueguchi Tanaka M;Mizutani M;Sakata K;Takatsuto S;Yoshida S;Tanaka H;Kitano H;Matsuoka M
Erect leaves caused by brassinosteroid deficiency increase biomass production and grain yield in rice
Nature Biotechnology, 2006, 24(1): 105-109

==Structured Information==
{{JaponicaGene|
GeneName = Os03g0227700|
Description = Cytochrome P450 family protein|
Version = NM_001055982.2 GI:297600576 GeneID:4332134|
Length = 6529 bp|
Definition = Oryza sativa Japonica Group Os03g0227700, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 3|Chromosome 3]]|
AP = Chromosome 3:6765109..6771637|
CDS = 6765109..6765374,6765543..6765867,6765971..6766123,6768937..6769188,6769337..6769429 ,6770222..6770300,6770398..6770504,6771392..6771637|
GCID = <gbrowseImage1>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</cdnaseq>|
AA = <aaseq>MAAMMASITSELLFFLPFILLALLTFYTTTVAKCHGGHWWRGGT TPAKRKRMNLPPGAAGWPLVGETFGYLRAHPATSVGRFMEQHIARYGKIYRSSLFGER TVVSADAGLNRYILQNEGRLFECSYPRSIGGILGKWSMLVLVGDPHREMRAISLNFLS SVRLRAVLLPEVERHTLLVLRAWPPSSTFSAQHQAKKFTFNLMAKNIMSMDPGEEETE RLRREYITFMKGVVSAPLNLPGTPYWKALKSRAAILGVIERKMEERVEKLSKEDASVE QDDLLGWALKQSNLSKEQILDLLLSLLFAGHETSSMALALAIFFLEGCPKAVQELREE HLGIARRQRLRGECKLSWEDYKEMVFTQCVINETLRLGNVVRFLHRKVIKDVHYKGYD IPSGWKILPVLAAVHLDSSLYEDPQRFNPWRWKSSGSSGGLAQSSSFMPYGGGTRLCA GSELAKLEMAVFLHHLVLNFRWELAEPDQAFVFPFVDFPKGLPIRVHRIAQDDEQE</aaseq>|
DNA = <dnaseqindica>1..266#435..759#863..1015#3829..4080#4229..4321#5114..5192#5290..5396#6284..6529#atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtcggtggtttcattactcggtacacgcacacagctatataagcacgcgcactgctcgcctcgccgtgaagcaagcttagctagtagctaggatatcgatcgatcggtaataataggctgacggcgagctggggggtggcgtgcggttgcgtgcgtgcgtgcatgcaggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaaggtacaacacacatcgtcctccattaagttgagcatgcatatatcaatcagctcgctctagcgatcaatatcaatgaaacgcgcgtttgcgtgcgtgcgtgcagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaaggtacgtactcgtagctactagctactccctccgtttcgaaatgtttgacgccgttgactttttatcacatatttgaccgttcgttttatttaaaaaaattaagtaattattaattattttcctatcatttgattcattgttaaatatatttttatgtaggcatataattttacatatctcacaaaagtttttaaataagacgaacggtcaaacatttgctaaaaagtcaacggtgtcaaatatttcgaaacggaggaagtagctagctaccaccccacccgtgcggattgatcgatcgatcgatgtccctctctgcctacacacaccctgcgccccggccgtacgcgcgcaaagtcgtgcgcgcatgagttttactccttccgtttcaaaaaaaaaacgacctagacaagtaatatgatgtaacctagaacaacgaatctatacaggagtatatccaaattcattgtcctagattacatcatatctttgtctatgttgccttttttaggaggagagagtactttacatatgagttatgaccggccggccaggttcctttcttcccctattccgtatgcatactataagcagctgcatataggcatatactactatacccagcttagctaaagcaaacaacttgcaacgacgatcatgtgcaggtccctgttgacagaggttagagatgtaaaccatttttgttgttttcattatctaaagagaaaacaagttactacgacgatcgagatgtacgcaaggggatcatatttatattggagaggagagagagatgatttgcgacttgtgagtgataagtgaatatatatactatatatttgtgtaggagtgacgatgacttccctgtcaatgttttgtttggtatgcacacggggggaaaagagtttagagaaaggaattggctagagagggccaggtttagtttctttaatttttgtttggtggttttgtgtgtagctagtccgcatgcatgatatatcatatatatgcatggcccatcattagatatatatacatgacggcagtgcacttattatatatatgtaggggcatatacaaggtgtatgtgtttagaattgtggatggtccaggaattgttgcagctagttctatactagcatagtgaattagtgatccatatcgagatcaggtccaatcggtcagctcgcacagcgggtcttttagtacgatttgttcggacaaattatcttgcctttgagataatcaacataggattgaaactgcatactagcaatagaccatagctatttggcaatatagaaagctaaaaaatgttgattgaaaggtgcgtgcatgcatgctagcagtatgtcttttaggccaggtccagtttttttttttttctcaaaggcagttgcttaaggtgaaattctagcagaggtcatatgaatatatttattatcttccaaacatgagccaaaacctatatatagttaactcaaaaaaccatgcattggtacttcttctgagtatgcatcctggggactgatcgatcatgcagattcttttctttctttttccctggggacgtactgatcatcactcatgtgagcataaaatgttgcctactttttactgcatgtgaggtcacactcaaatgatctttattgattgcctcagcatgcatgctgctagctttcacgatctctctccgatacacacaccctgtgctgtgctcaacagcatgcaacctggcaggtggaatttttaatctgttggaaagtaaatgaccaaccttcataatttggcatgattatatgctggggaatctatcatttatcatgattcaaccgctgaccaaactctttgccacacacccgaaccacattaatttcgttaagcggatgattagccagcaaccttcacagtttggttttaattagttaatgaaagggtgcttgtagttttattagctgaagagacactgcagtactcctactgagcgttctccttaattgggaaatagatctctgcagtactatcctttaactttaagtgaataacatgtagctaggtctcacacttatcaggcccacacatcagtgatccaagtcaaatggcaatacaccataggagattcttccactaattggctaacaaaaataaactgtgatcactgatcacagctgctaacactataacacattaatttgcatgtctcatgaaatgccagatgagttggattaattatctatcttatgttcagtgccgaggaatatacaaggaaacatcattggtatagttgtggagttctgagtgcattatctcacccagtacggtcaaattaatcctagtctccgtaaatatggttgtgtgcactcttagttaattagtgtagacccggaaattatactttgtcgagaatgttgttatccgtgtgttaagtttcaagtaccctgcaggcctataactaaacactgccaacatggatgtttaagaatgaattatacatacagcgcacttggaaaagaaaaggggtagctacccagctgaagaacacctctctctcatttgaaggtcgacacaatatgtgtcttttctatatcgaaaggaaacgacattttttctcttcaaaacagaaagctatgacaagagacataatcctagactagtaggcggttaggcaacaccatatctgaatttgataggtgtcccacatattttttttcttttcctgataagaaggaacttttcaccgtgagcttatttttttctgttctatgtcgacattagttccatgatgtttactgtgtatttttactgttcccctgcagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggtacagagaattacagattcgaaattaacatctactgtcactagctcagtgctacgtttaactgtagagcaacaattgcagaaagcgaaggcatcagaaatgacatgggaatcgattagttacactcacactgacgcgtgacatgcaggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtaaggccccccctactttgttccttccgatccctcctccctaaacgctactaatataaatcggtgcaaaggctttgtgctagcgtagctcttgctgttttgtttggtcacttgcgtagagagatagcaacttcccgaaatgtcatgtacgaatggagactgaaagctctgcgaaataattgccgggccacgtactacagtacttacgctccctctcttcagtttggatcgatcgagctaagaaggcgcacgcatgtgtgcattgatcgatagagtcgatctgttctgccttgtaagctagccagctgaggttctttcgccttggaagtttctacagggtgaggcccatcgtcttgcaacgatccaaatgccatgacaagtacctgttgttgtaggcttccagtagcacggcctgcccccaagaaggtatctgaatcttctgcattgctgttcagacggcagagcagaacatgccgaaataccattccagtcagttgcagtatatgtgtgtggtggtatcagataaaggagcagagtaatttgaggccttagaaattcagatgagttgctcgacaattgtcgcctaccgatcgagaataagttgcatgcatgtatgtgatgtaattatgggtcttagtcagcaggatagtttagtcgctgcagtactgtagattgtagtacttgcgttcaaagcagacccaaaacagaacttgcattcaggaaacaaaacatgaaaaaagaaatatacagtacttggcactgatatttcttttgtttgggtccttattgcaggtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggtgagtagctcgatttcactcttcttccctgaatcgacctctcctctatctgacagcagaagctaactgaactgaactgaactgaactgaactgaaggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaaggtcagtcgcagtaggattatcagtgaatctaatctattaatctcggtttaattgctgggattcaaggaaaagaaaatactgggacaagcgcggatgtcgctgttgggcagtgtctgtcaatcataagcgaaggggatcatgtcagaatctgcaaaagcagcacaggctaaacgcgtgcttgcctcgagcagcaatcatgcagagaggcaagctccgcgataggaacatgcatggtgaatcactgaacctgatatgaaagtgaatgctcagctcagctcgctcgctcgcgcgcgcatgtgctttctcatttcgccgtgttgcgtgctagctagtcaatggtgttcccacgaattgttttctcttctcatcacacaaagtcagtggtgaaaaccgaggtgcatgtgcggtaatggaacgggggattaggcggcgacgccgtgatggattatgcattcgtgcgagaaaaatccaggagcgttggtccaaatcgcaagcggatcaaaccgaaccaatcagtggacgattggagccctgcagtccatccatcacatctccattcttatccgctgcgactcacggacattagttgttgaggggtgaacgacgcatgtgtgtatgtatgctcgatcgatcgatcgaacaatggatctgtgtgtggtgtagttgccaacgccgacgagattatctgctcgtctgcatgcagccgcgaaaaagacatgtgtccggattatcctgcaggagattgagcaagataatctgctgcgcaaaaactgcagccaatcgatcattccatgacgcgaaaaaaaaaacaataattactactgctgatacgataacttcgcttcgaattgttgacacacaagtgttcttcttttgttcgattgcagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001055982.2 RefSeq:Os03g0227700]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 3]]
[[Category:Chromosome 3]]

Os03g0227700

2014-05-24T12:52:39Z

LQ: /* Annotated Information */

Please input one-sentence summary here.
''OsDWAR''F4 is expressed in young elongating tissues.
==Annotated Information==
Brassinosteroids influence both plant height and leaf erectness in rice[6]. Arabidopsis thaliana BRASSINOSTEROID INSENSITIVE1 (bri1) mutants, which are deficient in the brassinosteroid receptor, have a severe dwarf phenotype[7]. The rice mutants d61-1 and d61-2, which are weak mutant alleles of OsBRI1, a homolog of A. thaliana BRI1, have both a semi-dwarf phenotype and more erect leaves[8]. In rice, an erect leaf phenotype has also been associated with loss-of-function, brassinosteroid-deficient mutants such as brassinosteroid-deficient dwarf1 (brd1), which is defective in OsDWARF (the homolog of tomato CYP85A1/DWARF[9]), and ebisu dwarf (d2), which is defective in CYP90D2/D2 (ref. [10]). However, the grain yield of d61, brd1 and d2 mutants is decreased because of morphological alterations in their reproductive development.

===Function===
New cultivars with very erect leaves, which increase light capture for photosynthesis and nitrogen storage for grain filling, may have increased grain yields.The erect leaf phenotype of a rice brassinosteroid-deficient mutant, osdwarf4-1, is associated with enhanced grain yields under conditions of dense planting, even without extra fertilizer. Molecular and biochemical studies reveal that two different cytochrome P450s, CYP90B2/OsDWARF4 and CYP724B1/D11, function redundantly in C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis. Therefore, despite the central role of brassinosteroids in plant growth and development, mutation of OsDWARF4 alone causes only limited defects in brassinosteroid biosynthesis and plant morphology[1].

===Expression===
The affected gene, OsDWARF4, was identified by characterizing the site of insertion of the Tos17 retrotransposon.

----Molecular characterization of OsDWARF4 and OsDWARF4L1/D11
OsDWARF4 is a homolog of A. thaliana DWARF4, which encodes a cytochrome P450, CYP90B1 (Fig. 1a). CYP90B1/DWARF4 catalyzes C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis[11]. Only one DWARF4 homolog was identified in the rice genome. OsDWARF4 is located on the short arm of chromosome 3 (35 cM). Its predicted open reading frame consists of eight exons, and encodes a protein of 502 amino acids (Fig. 1b,c). OsDWARF4 is most closely related to DWARF4 (66.3% amino-acid identity; Fig. 1a). The structure of OsDWARF4 is similar to that of DWARF4 throughout its length: five domains found in cytochrome P450s—proline rich, A (also referred to as dioxygen binding), B (steroid binding), C and heme binding—are highly conserved (Fig. 1c). In wild-type rice, OsDWARF4 transcripts are most abundant in leaf blades and roots, but are also found in all other plant parts tested (Fig. 1d). Transcript levels were decreased by exogenous brassinolide treatment, and increased in brassinosteroid-insensitive d61-3 and brassinosteroid-deficient brd1-1 (Fig. 1e). These results suggest that OsDWARF4 expression is feedback regulated, as is that of A. thaliana brassinosteroid-biosynthesis genes[12].
OsDWARF4L1 was identified as an OsDWARF4-like gene by in silico screening of rice DNA databases. OsDWARF4L1 is located on the long arm of chromosome 4 (70.9 cM). A recent study demonstrated that a rice semi-dwarf and short-grain mutant, d11, has a loss-of-function mutation in the OsDWARF4L1 gene13. OsDWARF4L1/D11 shows the highest homology with A. thaliana CYP724A1, and is closely related to rice OsDWARF4 and A. thaliana DWARF4 (47.4%, 37.7% and 41.7% amino-acid identities, respectively; Fig. 1a). In wild-type rice, OsDWARF4L1/D11 transcripts were most abundant in vegetative shoot apices, leaf sheaths and roots, and found at more moderate levels in other organs (Fig. 1d). They were decreased by exogenous brassinolide treatment, and increased in d61-3 and brd1-1 (Fig. 1e). These results indicate that OsDWARF4L1/D11 expression is regulated by a feedback mechanism, as is that of OsDWARF4.

----Phenotypes of rice mutants defective in brassinosteroid biosynthesis
In osdwarf4-1, Tos17 insertion into exon 6 causes a premature termination in domain C (arrowhead in Fig. 1b,c). Because a premature stop codon before the heme-binding domain caused loss-of-function of A. thaliana DWARF4 (ref. 11), we consider osdwarf4-1 to be a null allele. Unexpectedly, osdwarf4-1 plants show only weak morphological phenotypes, such as a slightly dwarfed stature and more erect leaves (Fig. 2a,b). Both of these phenotypes were complemented by the introduction of an entire OsDWARF4 gene (data not shown). These results seem inconsistent with the fact that rice d61-3 and brd1-1, which are loss-of-function mutants of OsBRI1 and OsDWARF respectively, display severe dwarfing and have malformed leaves (for example, see brd1-1 in Fig. 2a), and suggest functional redundancy in brassinosteroid biosynthesis in rice.
Because null alleles of d11, such as d11-4, also show a weak phenotype13 (Fig. 2a), we suspected that OsDWARF4L1/D11 and OsDWARF4 may function redundantly. To confirm this possibility, we produced double mutants by crossing homozygous osdwarf4-1 and d11-4 plants. Among the F2 population, 34 of 576 plants showed severe dwarfing and malformed leaves with tortuous leaf blades, indistinguishable from the brassinosteroid-deficient brd1-1 phenotype (Fig. 2a,c,d). This ratio fits the theoretical 9:3:3:1 (wild-type:osdwarf4-1:d11-4:osdwarf4-1/d11-4) segregation predicted by mendelian inheritance. PCR and sequence analyses confirmed that these plants were osdwarf4-1/d11-4 double mutants (data not shown).
Endogenous levels of brassinosteroid intermediates in brd1 clearly indicate that OsDWARF catalyzes C-6 oxidation9. However, quantification in d2 could not determine the step(s) catalyzed by CYP90D2/D2, because CYP90D2/D2 functions redundantly with another gene, CYP90D3, and loss-of-function of CYP90D2/D2 have caused only limited defects in brassinosteroid biosynthesis10. Similarly, endogenous levels of brassinosteroid intermediates were obviously changed in the osdwarf4-1/d11-4 double mutant relative to osdwarf4-1 or d11 single mutants. Although brassinolide was not detected in either the wild-type or osdwarf4-1/d11-4, another bioactive brassinosteroid, castasterone (CS), was detected in the wild-type but not in osdwarf4-1/d11-4, confirming that osdwarf4-1/d11-4 is brassinosteroid-deficient (Fig. 2e). Levels of the other five intermediates downstream of C-22 hydroxylation—6-deoxocathasterone (6-DeoxoCT), 6-deoxoteasterone (6-DeoxoTE), 3-dehydro-6-deoxoteasterone (6-Deoxo3DT), 6-deoxotyphasterol (6-DeoxoTY) and 6-deoxocastasterone (6-DeoxoCS)—were also greatly reduced in osdwarf4-1/d11-4 (Fig. 2e). This indicates that both proteins are involved in C-22 hydroxylation. It is unclear why the upstream compound, campestanol, did not accumulate in osdwarf4-1/d11-4. Because A. thaliana CYP90B1/DWARF4 also catalyzes C-22 hydroxylation of other upstream compounds such as campesterol and (24R)-ergost-4-en-3-one14, it is possible that campestanol is not the major substrate for CYP90B2/OsDWARF4 and CYP724B1/D11 in rice.

----Characterization of a field-grown osdwarf4-1 mutant
Because the harvest index (grain/(grain plus straw)) of rice is about 0.5, further increases in yield potential will have to involve increases in crop biomass driven by more net photosynthesis15. In rice, the contribution of lower leaves to photosynthesis is significant, even though their photosynthetic capacity is considerably less than that of upper leaves16. Erect leaves allow greater penetration of light to lower leaves, thereby optimizing canopy photosynthesis1. Therefore, the most likely explanation for the increase in above-ground biomass in osdwarf4-1 plots under dense planting conditions (more than 1.2 times that in wild-type under conventional planting condition) is that the shade of the upper leaves is minimized, and the lower leaves receive more light to drive higher rates of photosynthesis (Fig. 3 and Supplementary Table 2 online). No difference was observed in the heading date (flowering time) and the grain-filling period (time to maturity) between wild-type and osdwarf4-1 plants (data not shown). In the wild-type, the above-ground biomass was increased in response to higher fertilizer application under conventional planting (22.2 plants m-2), but was significantly decreased under dense planting with doubled fertilizer application owing to reduced photosynthesis caused by lodging (Fig. 3a). In contrast, the above-ground biomass of osdwarf4-1 was significantly (P < 0.01) increased by dense planting at each fertilizer rate. Consequently, the above-ground biomass in osdwarf4-1 plots under dense planting with standard and a 1.5-fold increase in fertilizer application was approx1.34 and 1.52 times that in wild-type plots under conventional cultivation conditions. The increased biomass depended mainly on accelerated formation of tillers, which have the ability to generate a panicle (panicle number, Fig. 3b). Both total and fertile grain numbers tended to increase in densely planted osdwarf4-1 plots (Fig. 3c), resulting in significantly increased grain yield even when the fertilizer application was not increased (for example, the osdwarf4-1 yield was 6.19 t ha-1 under dense planting with standard fertilizer application whereas the wild-type yield was 4.69 t ha-1 under conventional conditions; Fig. 3d). However, the increase in grain yield was not as much as that in above-ground biomass, because the harvest index of osdwarf4-1 (0.41) was slightly lower than that of the wild-type (0.44). A similar field test gave comparable results.

----Overexpression of OsDWARF4
Overexpression of DWARF in transgenic tomato increases plant height17, and overexpression of DWARF4 in transgenic A. thaliana and tobacco increases vegetative growth and seed yield18. In rice, however, the role of brassinosteroid in regulating leaf angle has been known for over 40 years19, 20. Whereas brassinosteroid deficiency increased the erectness of leaves, the opposite phenotype (increased leaf angle) was observed in transgenic rice overexpressing OsDWARF4 (27.6° versus 10.0° in the wild-type, P < 0.01), and was inherited in their progeny (data not shown). Because inclined leaves are unfavorable for photosynthesis when plants are grown in a group, we have to carefully evaluate whether the overexpression of a brassinosteroid-biosynthesis gene in transgenic rice increases grain yield as is the case in transgenic plants overexpressing DWARF4.

===Conclusion===
A previous study concluded that more erect leaves improve not only light distribution but also the size and use of leaf nitrogen reservoirs for grain growth1. Our results clearly demonstrate that the erect leaf phenotype caused by brassinosteroid deficiency improved biomass production under field conditions, resulting in increased grain yield without the negative environmental effects associated with fertilizers. We have recently identified rice homologs of A. thaliana BAS1 that encodes the brassinosteroid catabolic enzyme, CYP734A1 (ref. 21). Overexpression of these rice homologs in transgenic rice successfully induced the brassinosteroid-deficient phenotypes (unpublished results). Although ideally, expression of these genes should be restricted to particular tissues and organs such as leaf laminae, it should be possible to generate erect leaf varieties by the introduction of a single transgene. In conclusion, engineering more erect leaves modulating brassinosteroid metabolism can be combined with dense planting to improve rice grain yields.

==Labs working on this gene==
Please input related labs here.

==References==
1. Sakamoto T;Morinaka Y;Ohnishi T;Sunohara H;Fujioka S;Ueguchi Tanaka M;Mizutani M;Sakata K;Takatsuto S;Yoshida S;Tanaka H;Kitano H;Matsuoka M
Erect leaves caused by brassinosteroid deficiency increase biomass production and grain yield in rice
Nature Biotechnology, 2006, 24(1): 105-109

==Structured Information==
{{JaponicaGene|
GeneName = Os03g0227700|
Description = Cytochrome P450 family protein|
Version = NM_001055982.2 GI:297600576 GeneID:4332134|
Length = 6529 bp|
Definition = Oryza sativa Japonica Group Os03g0227700, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 3|Chromosome 3]]|
AP = Chromosome 3:6765109..6771637|
CDS = 6765109..6765374,6765543..6765867,6765971..6766123,6768937..6769188,6769337..6769429 ,6770222..6770300,6770398..6770504,6771392..6771637|
GCID = <gbrowseImage1>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</cdnaseq>|
AA = <aaseq>MAAMMASITSELLFFLPFILLALLTFYTTTVAKCHGGHWWRGGT TPAKRKRMNLPPGAAGWPLVGETFGYLRAHPATSVGRFMEQHIARYGKIYRSSLFGER TVVSADAGLNRYILQNEGRLFECSYPRSIGGILGKWSMLVLVGDPHREMRAISLNFLS SVRLRAVLLPEVERHTLLVLRAWPPSSTFSAQHQAKKFTFNLMAKNIMSMDPGEEETE RLRREYITFMKGVVSAPLNLPGTPYWKALKSRAAILGVIERKMEERVEKLSKEDASVE QDDLLGWALKQSNLSKEQILDLLLSLLFAGHETSSMALALAIFFLEGCPKAVQELREE HLGIARRQRLRGECKLSWEDYKEMVFTQCVINETLRLGNVVRFLHRKVIKDVHYKGYD IPSGWKILPVLAAVHLDSSLYEDPQRFNPWRWKSSGSSGGLAQSSSFMPYGGGTRLCA GSELAKLEMAVFLHHLVLNFRWELAEPDQAFVFPFVDFPKGLPIRVHRIAQDDEQE</aaseq>|
DNA = <dnaseqindica>1..266#435..759#863..1015#3829..4080#4229..4321#5114..5192#5290..5396#6284..6529#atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtcggtggtttcattactcggtacacgcacacagctatataagcacgcgcactgctcgcctcgccgtgaagcaagcttagctagtagctaggatatcgatcgatcggtaataataggctgacggcgagctggggggtggcgtgcggttgcgtgcgtgcgtgcatgcaggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaaggtacaacacacatcgtcctccattaagttgagcatgcatatatcaatcagctcgctctagcgatcaatatcaatgaaacgcgcgtttgcgtgcgtgcgtgcagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaaggtacgtactcgtagctactagctactccctccgtttcgaaatgtttgacgccgttgactttttatcacatatttgaccgttcgttttatttaaaaaaattaagtaattattaattattttcctatcatttgattcattgttaaatatatttttatgtaggcatataattttacatatctcacaaaagtttttaaataagacgaacggtcaaacatttgctaaaaagtcaacggtgtcaaatatttcgaaacggaggaagtagctagctaccaccccacccgtgcggattgatcgatcgatcgatgtccctctctgcctacacacaccctgcgccccggccgtacgcgcgcaaagtcgtgcgcgcatgagttttactccttccgtttcaaaaaaaaaacgacctagacaagtaatatgatgtaacctagaacaacgaatctatacaggagtatatccaaattcattgtcctagattacatcatatctttgtctatgttgccttttttaggaggagagagtactttacatatgagttatgaccggccggccaggttcctttcttcccctattccgtatgcatactataagcagctgcatataggcatatactactatacccagcttagctaaagcaaacaacttgcaacgacgatcatgtgcaggtccctgttgacagaggttagagatgtaaaccatttttgttgttttcattatctaaagagaaaacaagttactacgacgatcgagatgtacgcaaggggatcatatttatattggagaggagagagagatgatttgcgacttgtgagtgataagtgaatatatatactatatatttgtgtaggagtgacgatgacttccctgtcaatgttttgtttggtatgcacacggggggaaaagagtttagagaaaggaattggctagagagggccaggtttagtttctttaatttttgtttggtggttttgtgtgtagctagtccgcatgcatgatatatcatatatatgcatggcccatcattagatatatatacatgacggcagtgcacttattatatatatgtaggggcatatacaaggtgtatgtgtttagaattgtggatggtccaggaattgttgcagctagttctatactagcatagtgaattagtgatccatatcgagatcaggtccaatcggtcagctcgcacagcgggtcttttagtacgatttgttcggacaaattatcttgcctttgagataatcaacataggattgaaactgcatactagcaatagaccatagctatttggcaatatagaaagctaaaaaatgttgattgaaaggtgcgtgcatgcatgctagcagtatgtcttttaggccaggtccagtttttttttttttctcaaaggcagttgcttaaggtgaaattctagcagaggtcatatgaatatatttattatcttccaaacatgagccaaaacctatatatagttaactcaaaaaaccatgcattggtacttcttctgagtatgcatcctggggactgatcgatcatgcagattcttttctttctttttccctggggacgtactgatcatcactcatgtgagcataaaatgttgcctactttttactgcatgtgaggtcacactcaaatgatctttattgattgcctcagcatgcatgctgctagctttcacgatctctctccgatacacacaccctgtgctgtgctcaacagcatgcaacctggcaggtggaatttttaatctgttggaaagtaaatgaccaaccttcataatttggcatgattatatgctggggaatctatcatttatcatgattcaaccgctgaccaaactctttgccacacacccgaaccacattaatttcgttaagcggatgattagccagcaaccttcacagtttggttttaattagttaatgaaagggtgcttgtagttttattagctgaagagacactgcagtactcctactgagcgttctccttaattgggaaatagatctctgcagtactatcctttaactttaagtgaataacatgtagctaggtctcacacttatcaggcccacacatcagtgatccaagtcaaatggcaatacaccataggagattcttccactaattggctaacaaaaataaactgtgatcactgatcacagctgctaacactataacacattaatttgcatgtctcatgaaatgccagatgagttggattaattatctatcttatgttcagtgccgaggaatatacaaggaaacatcattggtatagttgtggagttctgagtgcattatctcacccagtacggtcaaattaatcctagtctccgtaaatatggttgtgtgcactcttagttaattagtgtagacccggaaattatactttgtcgagaatgttgttatccgtgtgttaagtttcaagtaccctgcaggcctataactaaacactgccaacatggatgtttaagaatgaattatacatacagcgcacttggaaaagaaaaggggtagctacccagctgaagaacacctctctctcatttgaaggtcgacacaatatgtgtcttttctatatcgaaaggaaacgacattttttctcttcaaaacagaaagctatgacaagagacataatcctagactagtaggcggttaggcaacaccatatctgaatttgataggtgtcccacatattttttttcttttcctgataagaaggaacttttcaccgtgagcttatttttttctgttctatgtcgacattagttccatgatgtttactgtgtatttttactgttcccctgcagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggtacagagaattacagattcgaaattaacatctactgtcactagctcagtgctacgtttaactgtagagcaacaattgcagaaagcgaaggcatcagaaatgacatgggaatcgattagttacactcacactgacgcgtgacatgcaggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtaaggccccccctactttgttccttccgatccctcctccctaaacgctactaatataaatcggtgcaaaggctttgtgctagcgtagctcttgctgttttgtttggtcacttgcgtagagagatagcaacttcccgaaatgtcatgtacgaatggagactgaaagctctgcgaaataattgccgggccacgtactacagtacttacgctccctctcttcagtttggatcgatcgagctaagaaggcgcacgcatgtgtgcattgatcgatagagtcgatctgttctgccttgtaagctagccagctgaggttctttcgccttggaagtttctacagggtgaggcccatcgtcttgcaacgatccaaatgccatgacaagtacctgttgttgtaggcttccagtagcacggcctgcccccaagaaggtatctgaatcttctgcattgctgttcagacggcagagcagaacatgccgaaataccattccagtcagttgcagtatatgtgtgtggtggtatcagataaaggagcagagtaatttgaggccttagaaattcagatgagttgctcgacaattgtcgcctaccgatcgagaataagttgcatgcatgtatgtgatgtaattatgggtcttagtcagcaggatagtttagtcgctgcagtactgtagattgtagtacttgcgttcaaagcagacccaaaacagaacttgcattcaggaaacaaaacatgaaaaaagaaatatacagtacttggcactgatatttcttttgtttgggtccttattgcaggtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggtgagtagctcgatttcactcttcttccctgaatcgacctctcctctatctgacagcagaagctaactgaactgaactgaactgaactgaactgaaggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaaggtcagtcgcagtaggattatcagtgaatctaatctattaatctcggtttaattgctgggattcaaggaaaagaaaatactgggacaagcgcggatgtcgctgttgggcagtgtctgtcaatcataagcgaaggggatcatgtcagaatctgcaaaagcagcacaggctaaacgcgtgcttgcctcgagcagcaatcatgcagagaggcaagctccgcgataggaacatgcatggtgaatcactgaacctgatatgaaagtgaatgctcagctcagctcgctcgctcgcgcgcgcatgtgctttctcatttcgccgtgttgcgtgctagctagtcaatggtgttcccacgaattgttttctcttctcatcacacaaagtcagtggtgaaaaccgaggtgcatgtgcggtaatggaacgggggattaggcggcgacgccgtgatggattatgcattcgtgcgagaaaaatccaggagcgttggtccaaatcgcaagcggatcaaaccgaaccaatcagtggacgattggagccctgcagtccatccatcacatctccattcttatccgctgcgactcacggacattagttgttgaggggtgaacgacgcatgtgtgtatgtatgctcgatcgatcgatcgaacaatggatctgtgtgtggtgtagttgccaacgccgacgagattatctgctcgtctgcatgcagccgcgaaaaagacatgtgtccggattatcctgcaggagattgagcaagataatctgctgcgcaaaaactgcagccaatcgatcattccatgacgcgaaaaaaaaaacaataattactactgctgatacgataacttcgcttcgaattgttgacacacaagtgttcttcttttgttcgattgcagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001055982.2 RefSeq:Os03g0227700]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 3]]
[[Category:Chromosome 3]]

Os03g0227700

2014-05-24T11:55:29Z

LQ: /* Annotated Information */

Please input one-sentence summary here.
''OsDWAR''F4 is expressed in young elongating tissues.
==Annotated Information==
OsDWARF4 is a homolog of A. thaliana DWARF4, which encodes a cytochrome P450, CYP90B1 (Fig. 1a). CYP90B1/DWARF4 catalyzes C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis[11]. Only one DWARF4 homolog was identified in the rice genome. OsDWARF4 is located on the short arm of chromosome 3 (35 cM). Its predicted open reading frame consists of eight exons, and encodes a protein of 502 amino acids (Fig. 1b,c). OsDWARF4 is most closely related to DWARF4 (66.3% amino-acid identity; Fig. 1a). The structure of OsDWARF4 is similar to that of DWARF4 throughout its length: five domains found in cytochrome P450s—proline rich, A (also referred to as dioxygen binding), B (steroid binding), C and heme binding—are highly conserved (Fig. 1c). In wild-type rice, OsDWARF4 transcripts are most abundant in leaf blades and roots, but are also found in all other plant parts tested (Fig. 1d). Transcript levels were decreased by exogenous brassinolide treatment, and increased in brassinosteroid-insensitive d61-3 and brassinosteroid-deficient brd1-1 (Fig. 1e). These results suggest that OsDWARF4 expression is feedback regulated, as is that of A. thaliana brassinosteroid-biosynthesis genes[12].

===Function===
New cultivars with very erect leaves, which increase light capture for photosynthesis and nitrogen storage for grain filling, may have increased grain yields.The erect leaf phenotype of a rice brassinosteroid-deficient mutant, osdwarf4-1, is associated with enhanced grain yields under conditions of dense planting, even without extra fertilizer. Molecular and biochemical studies reveal that two different cytochrome P450s, CYP90B2/OsDWARF4 and CYP724B1/D11, function redundantly in C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis. Therefore, despite the central role of brassinosteroids in plant growth and development, mutation of OsDWARF4 alone causes only limited defects in brassinosteroid biosynthesis and plant morphology[1].

===Expression===
Brassinosteroids influence both plant height and leaf erectness in rice[6]. Arabidopsis thaliana BRASSINOSTEROID INSENSITIVE1 (bri1) mutants, which are deficient in the brassinosteroid receptor, have a severe dwarf phenotype[7]. The rice mutants d61-1 and d61-2, which are weak mutant alleles of OsBRI1, a homolog of A. thaliana BRI1, have both a semi-dwarf phenotype and more erect leaves[8]. In rice, an erect leaf phenotype has also been associated with loss-of-function, brassinosteroid-deficient mutants such as brassinosteroid-deficient dwarf1 (brd1), which is defective in OsDWARF (the homolog of tomato CYP85A1/DWARF[9]), and ebisu dwarf (d2), which is defective in CYP90D2/D2 (ref. [10]). However, the grain yield of d61, brd1 and d2 mutants is decreased because of morphological alterations in their reproductive development.

===Evolution===
Please input evolution information here.

You can also add sub-section(s) at will.

==Labs working on this gene==
Please input related labs here.

==References==
1. Sakamoto T;Morinaka Y;Ohnishi T;Sunohara H;Fujioka S;Ueguchi Tanaka M;Mizutani M;Sakata K;Takatsuto S;Yoshida S;Tanaka H;Kitano H;Matsuoka M
Erect leaves caused by brassinosteroid deficiency increase biomass production and grain yield in rice
Nature Biotechnology, 2006, 24(1): 105-109

==Structured Information==
{{JaponicaGene|
GeneName = Os03g0227700|
Description = Cytochrome P450 family protein|
Version = NM_001055982.2 GI:297600576 GeneID:4332134|
Length = 6529 bp|
Definition = Oryza sativa Japonica Group Os03g0227700, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 3|Chromosome 3]]|
AP = Chromosome 3:6765109..6771637|
CDS = 6765109..6765374,6765543..6765867,6765971..6766123,6768937..6769188,6769337..6769429 ,6770222..6770300,6770398..6770504,6771392..6771637|
GCID = <gbrowseImage1>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</cdnaseq>|
AA = <aaseq>MAAMMASITSELLFFLPFILLALLTFYTTTVAKCHGGHWWRGGT TPAKRKRMNLPPGAAGWPLVGETFGYLRAHPATSVGRFMEQHIARYGKIYRSSLFGER TVVSADAGLNRYILQNEGRLFECSYPRSIGGILGKWSMLVLVGDPHREMRAISLNFLS SVRLRAVLLPEVERHTLLVLRAWPPSSTFSAQHQAKKFTFNLMAKNIMSMDPGEEETE RLRREYITFMKGVVSAPLNLPGTPYWKALKSRAAILGVIERKMEERVEKLSKEDASVE QDDLLGWALKQSNLSKEQILDLLLSLLFAGHETSSMALALAIFFLEGCPKAVQELREE HLGIARRQRLRGECKLSWEDYKEMVFTQCVINETLRLGNVVRFLHRKVIKDVHYKGYD IPSGWKILPVLAAVHLDSSLYEDPQRFNPWRWKSSGSSGGLAQSSSFMPYGGGTRLCA GSELAKLEMAVFLHHLVLNFRWELAEPDQAFVFPFVDFPKGLPIRVHRIAQDDEQE</aaseq>|
DNA = <dnaseqindica>1..266#435..759#863..1015#3829..4080#4229..4321#5114..5192#5290..5396#6284..6529#atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtcggtggtttcattactcggtacacgcacacagctatataagcacgcgcactgctcgcctcgccgtgaagcaagcttagctagtagctaggatatcgatcgatcggtaataataggctgacggcgagctggggggtggcgtgcggttgcgtgcgtgcgtgcatgcaggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaaggtacaacacacatcgtcctccattaagttgagcatgcatatatcaatcagctcgctctagcgatcaatatcaatgaaacgcgcgtttgcgtgcgtgcgtgcagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaaggtacgtactcgtagctactagctactccctccgtttcgaaatgtttgacgccgttgactttttatcacatatttgaccgttcgttttatttaaaaaaattaagtaattattaattattttcctatcatttgattcattgttaaatatatttttatgtaggcatataattttacatatctcacaaaagtttttaaataagacgaacggtcaaacatttgctaaaaagtcaacggtgtcaaatatttcgaaacggaggaagtagctagctaccaccccacccgtgcggattgatcgatcgatcgatgtccctctctgcctacacacaccctgcgccccggccgtacgcgcgcaaagtcgtgcgcgcatgagttttactccttccgtttcaaaaaaaaaacgacctagacaagtaatatgatgtaacctagaacaacgaatctatacaggagtatatccaaattcattgtcctagattacatcatatctttgtctatgttgccttttttaggaggagagagtactttacatatgagttatgaccggccggccaggttcctttcttcccctattccgtatgcatactataagcagctgcatataggcatatactactatacccagcttagctaaagcaaacaacttgcaacgacgatcatgtgcaggtccctgttgacagaggttagagatgtaaaccatttttgttgttttcattatctaaagagaaaacaagttactacgacgatcgagatgtacgcaaggggatcatatttatattggagaggagagagagatgatttgcgacttgtgagtgataagtgaatatatatactatatatttgtgtaggagtgacgatgacttccctgtcaatgttttgtttggtatgcacacggggggaaaagagtttagagaaaggaattggctagagagggccaggtttagtttctttaatttttgtttggtggttttgtgtgtagctagtccgcatgcatgatatatcatatatatgcatggcccatcattagatatatatacatgacggcagtgcacttattatatatatgtaggggcatatacaaggtgtatgtgtttagaattgtggatggtccaggaattgttgcagctagttctatactagcatagtgaattagtgatccatatcgagatcaggtccaatcggtcagctcgcacagcgggtcttttagtacgatttgttcggacaaattatcttgcctttgagataatcaacataggattgaaactgcatactagcaatagaccatagctatttggcaatatagaaagctaaaaaatgttgattgaaaggtgcgtgcatgcatgctagcagtatgtcttttaggccaggtccagtttttttttttttctcaaaggcagttgcttaaggtgaaattctagcagaggtcatatgaatatatttattatcttccaaacatgagccaaaacctatatatagttaactcaaaaaaccatgcattggtacttcttctgagtatgcatcctggggactgatcgatcatgcagattcttttctttctttttccctggggacgtactgatcatcactcatgtgagcataaaatgttgcctactttttactgcatgtgaggtcacactcaaatgatctttattgattgcctcagcatgcatgctgctagctttcacgatctctctccgatacacacaccctgtgctgtgctcaacagcatgcaacctggcaggtggaatttttaatctgttggaaagtaaatgaccaaccttcataatttggcatgattatatgctggggaatctatcatttatcatgattcaaccgctgaccaaactctttgccacacacccgaaccacattaatttcgttaagcggatgattagccagcaaccttcacagtttggttttaattagttaatgaaagggtgcttgtagttttattagctgaagagacactgcagtactcctactgagcgttctccttaattgggaaatagatctctgcagtactatcctttaactttaagtgaataacatgtagctaggtctcacacttatcaggcccacacatcagtgatccaagtcaaatggcaatacaccataggagattcttccactaattggctaacaaaaataaactgtgatcactgatcacagctgctaacactataacacattaatttgcatgtctcatgaaatgccagatgagttggattaattatctatcttatgttcagtgccgaggaatatacaaggaaacatcattggtatagttgtggagttctgagtgcattatctcacccagtacggtcaaattaatcctagtctccgtaaatatggttgtgtgcactcttagttaattagtgtagacccggaaattatactttgtcgagaatgttgttatccgtgtgttaagtttcaagtaccctgcaggcctataactaaacactgccaacatggatgtttaagaatgaattatacatacagcgcacttggaaaagaaaaggggtagctacccagctgaagaacacctctctctcatttgaaggtcgacacaatatgtgtcttttctatatcgaaaggaaacgacattttttctcttcaaaacagaaagctatgacaagagacataatcctagactagtaggcggttaggcaacaccatatctgaatttgataggtgtcccacatattttttttcttttcctgataagaaggaacttttcaccgtgagcttatttttttctgttctatgtcgacattagttccatgatgtttactgtgtatttttactgttcccctgcagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggtacagagaattacagattcgaaattaacatctactgtcactagctcagtgctacgtttaactgtagagcaacaattgcagaaagcgaaggcatcagaaatgacatgggaatcgattagttacactcacactgacgcgtgacatgcaggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtaaggccccccctactttgttccttccgatccctcctccctaaacgctactaatataaatcggtgcaaaggctttgtgctagcgtagctcttgctgttttgtttggtcacttgcgtagagagatagcaacttcccgaaatgtcatgtacgaatggagactgaaagctctgcgaaataattgccgggccacgtactacagtacttacgctccctctcttcagtttggatcgatcgagctaagaaggcgcacgcatgtgtgcattgatcgatagagtcgatctgttctgccttgtaagctagccagctgaggttctttcgccttggaagtttctacagggtgaggcccatcgtcttgcaacgatccaaatgccatgacaagtacctgttgttgtaggcttccagtagcacggcctgcccccaagaaggtatctgaatcttctgcattgctgttcagacggcagagcagaacatgccgaaataccattccagtcagttgcagtatatgtgtgtggtggtatcagataaaggagcagagtaatttgaggccttagaaattcagatgagttgctcgacaattgtcgcctaccgatcgagaataagttgcatgcatgtatgtgatgtaattatgggtcttagtcagcaggatagtttagtcgctgcagtactgtagattgtagtacttgcgttcaaagcagacccaaaacagaacttgcattcaggaaacaaaacatgaaaaaagaaatatacagtacttggcactgatatttcttttgtttgggtccttattgcaggtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggtgagtagctcgatttcactcttcttccctgaatcgacctctcctctatctgacagcagaagctaactgaactgaactgaactgaactgaactgaaggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaaggtcagtcgcagtaggattatcagtgaatctaatctattaatctcggtttaattgctgggattcaaggaaaagaaaatactgggacaagcgcggatgtcgctgttgggcagtgtctgtcaatcataagcgaaggggatcatgtcagaatctgcaaaagcagcacaggctaaacgcgtgcttgcctcgagcagcaatcatgcagagaggcaagctccgcgataggaacatgcatggtgaatcactgaacctgatatgaaagtgaatgctcagctcagctcgctcgctcgcgcgcgcatgtgctttctcatttcgccgtgttgcgtgctagctagtcaatggtgttcccacgaattgttttctcttctcatcacacaaagtcagtggtgaaaaccgaggtgcatgtgcggtaatggaacgggggattaggcggcgacgccgtgatggattatgcattcgtgcgagaaaaatccaggagcgttggtccaaatcgcaagcggatcaaaccgaaccaatcagtggacgattggagccctgcagtccatccatcacatctccattcttatccgctgcgactcacggacattagttgttgaggggtgaacgacgcatgtgtgtatgtatgctcgatcgatcgatcgaacaatggatctgtgtgtggtgtagttgccaacgccgacgagattatctgctcgtctgcatgcagccgcgaaaaagacatgtgtccggattatcctgcaggagattgagcaagataatctgctgcgcaaaaactgcagccaatcgatcattccatgacgcgaaaaaaaaaacaataattactactgctgatacgataacttcgcttcgaattgttgacacacaagtgttcttcttttgttcgattgcagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001055982.2 RefSeq:Os03g0227700]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 3]]
[[Category:Chromosome 3]]

Os03g0227700

2014-05-24T11:41:18Z

LQ: /* Annotated Information */

Os03g0227700

2014-05-24T11:20:08Z

LQ: /* Annotated Information */

Please input one-sentence summary here.
''OsDWAR''F4 is expressed in young elongating tissues.
==Annotated Information==

===Function===
New cultivars with very erect leaves, which increase light capture for photosynthesis and nitrogen storage for grain filling, may have increased grain yields. Here we show that the erect leaf phenotype of a rice brassinosteroid-deficient mutant, osdwarf4-1, is associated with enhanced grain yields under conditions of dense planting, even without extra fertilizer. Molecular and biochemical studies reveal that two different cytochrome P450s, CYP90B2/OsDWARF4 and CYP724B1/D11, function redundantly in C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis. Therefore, despite the central role of brassinosteroids in plant growth and development, mutation of OsDWARF4 alone causes only limited defects in brassinosteroid biosynthesis and plant morphology[1].

===Expression===
Please input expression information here.

===Evolution===
Please input evolution information here.

You can also add sub-section(s) at will.

==Labs working on this gene==
Please input related labs here.

==References==
1. Sakamoto T;Morinaka Y;Ohnishi T;Sunohara H;Fujioka S;Ueguchi Tanaka M;Mizutani M;Sakata K;Takatsuto S;Yoshida S;Tanaka H;Kitano H;Matsuoka M
Erect leaves caused by brassinosteroid deficiency increase biomass production and grain yield in rice
Nature Biotechnology, 2006, 24(1): 105-109

==Structured Information==
{{JaponicaGene|
GeneName = Os03g0227700|
Description = Cytochrome P450 family protein|
Version = NM_001055982.2 GI:297600576 GeneID:4332134|
Length = 6529 bp|
Definition = Oryza sativa Japonica Group Os03g0227700, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 3|Chromosome 3]]|
AP = Chromosome 3:6765109..6771637|
CDS = 6765109..6765374,6765543..6765867,6765971..6766123,6768937..6769188,6769337..6769429 ,6770222..6770300,6770398..6770504,6771392..6771637|
GCID = <gbrowseImage1>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</cdnaseq>|
AA = <aaseq>MAAMMASITSELLFFLPFILLALLTFYTTTVAKCHGGHWWRGGT TPAKRKRMNLPPGAAGWPLVGETFGYLRAHPATSVGRFMEQHIARYGKIYRSSLFGER TVVSADAGLNRYILQNEGRLFECSYPRSIGGILGKWSMLVLVGDPHREMRAISLNFLS SVRLRAVLLPEVERHTLLVLRAWPPSSTFSAQHQAKKFTFNLMAKNIMSMDPGEEETE RLRREYITFMKGVVSAPLNLPGTPYWKALKSRAAILGVIERKMEERVEKLSKEDASVE QDDLLGWALKQSNLSKEQILDLLLSLLFAGHETSSMALALAIFFLEGCPKAVQELREE HLGIARRQRLRGECKLSWEDYKEMVFTQCVINETLRLGNVVRFLHRKVIKDVHYKGYD IPSGWKILPVLAAVHLDSSLYEDPQRFNPWRWKSSGSSGGLAQSSSFMPYGGGTRLCA GSELAKLEMAVFLHHLVLNFRWELAEPDQAFVFPFVDFPKGLPIRVHRIAQDDEQE</aaseq>|
DNA = <dnaseqindica>1..266#435..759#863..1015#3829..4080#4229..4321#5114..5192#5290..5396#6284..6529#atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtcggtggtttcattactcggtacacgcacacagctatataagcacgcgcactgctcgcctcgccgtgaagcaagcttagctagtagctaggatatcgatcgatcggtaataataggctgacggcgagctggggggtggcgtgcggttgcgtgcgtgcgtgcatgcaggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaaggtacaacacacatcgtcctccattaagttgagcatgcatatatcaatcagctcgctctagcgatcaatatcaatgaaacgcgcgtttgcgtgcgtgcgtgcagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaaggtacgtactcgtagctactagctactccctccgtttcgaaatgtttgacgccgttgactttttatcacatatttgaccgttcgttttatttaaaaaaattaagtaattattaattattttcctatcatttgattcattgttaaatatatttttatgtaggcatataattttacatatctcacaaaagtttttaaataagacgaacggtcaaacatttgctaaaaagtcaacggtgtcaaatatttcgaaacggaggaagtagctagctaccaccccacccgtgcggattgatcgatcgatcgatgtccctctctgcctacacacaccctgcgccccggccgtacgcgcgcaaagtcgtgcgcgcatgagttttactccttccgtttcaaaaaaaaaacgacctagacaagtaatatgatgtaacctagaacaacgaatctatacaggagtatatccaaattcattgtcctagattacatcatatctttgtctatgttgccttttttaggaggagagagtactttacatatgagttatgaccggccggccaggttcctttcttcccctattccgtatgcatactataagcagctgcatataggcatatactactatacccagcttagctaaagcaaacaacttgcaacgacgatcatgtgcaggtccctgttgacagaggttagagatgtaaaccatttttgttgttttcattatctaaagagaaaacaagttactacgacgatcgagatgtacgcaaggggatcatatttatattggagaggagagagagatgatttgcgacttgtgagtgataagtgaatatatatactatatatttgtgtaggagtgacgatgacttccctgtcaatgttttgtttggtatgcacacggggggaaaagagtttagagaaaggaattggctagagagggccaggtttagtttctttaatttttgtttggtggttttgtgtgtagctagtccgcatgcatgatatatcatatatatgcatggcccatcattagatatatatacatgacggcagtgcacttattatatatatgtaggggcatatacaaggtgtatgtgtttagaattgtggatggtccaggaattgttgcagctagttctatactagcatagtgaattagtgatccatatcgagatcaggtccaatcggtcagctcgcacagcgggtcttttagtacgatttgttcggacaaattatcttgcctttgagataatcaacataggattgaaactgcatactagcaatagaccatagctatttggcaatatagaaagctaaaaaatgttgattgaaaggtgcgtgcatgcatgctagcagtatgtcttttaggccaggtccagtttttttttttttctcaaaggcagttgcttaaggtgaaattctagcagaggtcatatgaatatatttattatcttccaaacatgagccaaaacctatatatagttaactcaaaaaaccatgcattggtacttcttctgagtatgcatcctggggactgatcgatcatgcagattcttttctttctttttccctggggacgtactgatcatcactcatgtgagcataaaatgttgcctactttttactgcatgtgaggtcacactcaaatgatctttattgattgcctcagcatgcatgctgctagctttcacgatctctctccgatacacacaccctgtgctgtgctcaacagcatgcaacctggcaggtggaatttttaatctgttggaaagtaaatgaccaaccttcataatttggcatgattatatgctggggaatctatcatttatcatgattcaaccgctgaccaaactctttgccacacacccgaaccacattaatttcgttaagcggatgattagccagcaaccttcacagtttggttttaattagttaatgaaagggtgcttgtagttttattagctgaagagacactgcagtactcctactgagcgttctccttaattgggaaatagatctctgcagtactatcctttaactttaagtgaataacatgtagctaggtctcacacttatcaggcccacacatcagtgatccaagtcaaatggcaatacaccataggagattcttccactaattggctaacaaaaataaactgtgatcactgatcacagctgctaacactataacacattaatttgcatgtctcatgaaatgccagatgagttggattaattatctatcttatgttcagtgccgaggaatatacaaggaaacatcattggtatagttgtggagttctgagtgcattatctcacccagtacggtcaaattaatcctagtctccgtaaatatggttgtgtgcactcttagttaattagtgtagacccggaaattatactttgtcgagaatgttgttatccgtgtgttaagtttcaagtaccctgcaggcctataactaaacactgccaacatggatgtttaagaatgaattatacatacagcgcacttggaaaagaaaaggggtagctacccagctgaagaacacctctctctcatttgaaggtcgacacaatatgtgtcttttctatatcgaaaggaaacgacattttttctcttcaaaacagaaagctatgacaagagacataatcctagactagtaggcggttaggcaacaccatatctgaatttgataggtgtcccacatattttttttcttttcctgataagaaggaacttttcaccgtgagcttatttttttctgttctatgtcgacattagttccatgatgtttactgtgtatttttactgttcccctgcagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggtacagagaattacagattcgaaattaacatctactgtcactagctcagtgctacgtttaactgtagagcaacaattgcagaaagcgaaggcatcagaaatgacatgggaatcgattagttacactcacactgacgcgtgacatgcaggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtaaggccccccctactttgttccttccgatccctcctccctaaacgctactaatataaatcggtgcaaaggctttgtgctagcgtagctcttgctgttttgtttggtcacttgcgtagagagatagcaacttcccgaaatgtcatgtacgaatggagactgaaagctctgcgaaataattgccgggccacgtactacagtacttacgctccctctcttcagtttggatcgatcgagctaagaaggcgcacgcatgtgtgcattgatcgatagagtcgatctgttctgccttgtaagctagccagctgaggttctttcgccttggaagtttctacagggtgaggcccatcgtcttgcaacgatccaaatgccatgacaagtacctgttgttgtaggcttccagtagcacggcctgcccccaagaaggtatctgaatcttctgcattgctgttcagacggcagagcagaacatgccgaaataccattccagtcagttgcagtatatgtgtgtggtggtatcagataaaggagcagagtaatttgaggccttagaaattcagatgagttgctcgacaattgtcgcctaccgatcgagaataagttgcatgcatgtatgtgatgtaattatgggtcttagtcagcaggatagtttagtcgctgcagtactgtagattgtagtacttgcgttcaaagcagacccaaaacagaacttgcattcaggaaacaaaacatgaaaaaagaaatatacagtacttggcactgatatttcttttgtttgggtccttattgcaggtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggtgagtagctcgatttcactcttcttccctgaatcgacctctcctctatctgacagcagaagctaactgaactgaactgaactgaactgaactgaaggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaaggtcagtcgcagtaggattatcagtgaatctaatctattaatctcggtttaattgctgggattcaaggaaaagaaaatactgggacaagcgcggatgtcgctgttgggcagtgtctgtcaatcataagcgaaggggatcatgtcagaatctgcaaaagcagcacaggctaaacgcgtgcttgcctcgagcagcaatcatgcagagaggcaagctccgcgataggaacatgcatggtgaatcactgaacctgatatgaaagtgaatgctcagctcagctcgctcgctcgcgcgcgcatgtgctttctcatttcgccgtgttgcgtgctagctagtcaatggtgttcccacgaattgttttctcttctcatcacacaaagtcagtggtgaaaaccgaggtgcatgtgcggtaatggaacgggggattaggcggcgacgccgtgatggattatgcattcgtgcgagaaaaatccaggagcgttggtccaaatcgcaagcggatcaaaccgaaccaatcagtggacgattggagccctgcagtccatccatcacatctccattcttatccgctgcgactcacggacattagttgttgaggggtgaacgacgcatgtgtgtatgtatgctcgatcgatcgatcgaacaatggatctgtgtgtggtgtagttgccaacgccgacgagattatctgctcgtctgcatgcagccgcgaaaaagacatgtgtccggattatcctgcaggagattgagcaagataatctgctgcgcaaaaactgcagccaatcgatcattccatgacgcgaaaaaaaaaacaataattactactgctgatacgataacttcgcttcgaattgttgacacacaagtgttcttcttttgttcgattgcagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001055982.2 RefSeq:Os03g0227700]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 3]]
[[Category:Chromosome 3]]

Os03g0227700

2014-05-24T11:12:27Z

LQ: /* Annotated Information */

Please input one-sentence summary here.
''OsDWAR''F4 is expressed in young elongating tissues.
==Annotated Information==
New cultivars with very erect leaves, which increase light capture for photosynthesis and nitrogen storage for grain filling, may have increased grain yields. Here we show that the erect leaf phenotype of a rice brassinosteroid-deficient mutant, osdwarf4-1, is associated with enhanced grain yields under conditions of dense planting, even without extra fertilizer. Molecular and biochemical studies reveal that two different cytochrome P450s, CYP90B2/OsDWARF4 and CYP724B1/D11, function redundantly in C-22 hydroxylation, the rate-limiting step of brassinosteroid biosynthesis. Therefore, despite the central role of brassinosteroids in plant growth and development, mutation of OsDWARF4 alone causes only limited defects in brassinosteroid biosynthesis and plant morphology. These results suggest that regulated genetic modulation of brassinosteroid biosynthesis can improve crops without the negative environmental effects of fertilizers[1].
===Function===
Please input function information here.
xxx

===Expression===
Please input expression information here.

===Evolution===
Please input evolution information here.

You can also add sub-section(s) at will.

==Labs working on this gene==
Please input related labs here.

==References==
1. Sakamoto T;Morinaka Y;Ohnishi T;Sunohara H;Fujioka S;Ueguchi Tanaka M;Mizutani M;Sakata K;Takatsuto S;Yoshida S;Tanaka H;Kitano H;Matsuoka M
Erect leaves caused by brassinosteroid deficiency increase biomass production and grain yield in rice
Nature Biotechnology, 2006, 24(1): 105-109

==Structured Information==
{{JaponicaGene|
GeneName = Os03g0227700|
Description = Cytochrome P450 family protein|
Version = NM_001055982.2 GI:297600576 GeneID:4332134|
Length = 6529 bp|
Definition = Oryza sativa Japonica Group Os03g0227700, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 3|Chromosome 3]]|
AP = Chromosome 3:6765109..6771637|
CDS = 6765109..6765374,6765543..6765867,6765971..6766123,6768937..6769188,6769337..6769429 ,6770222..6770300,6770398..6770504,6771392..6771637|
GCID = <gbrowseImage1>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008396:6765109..6771637
source=RiceChromosome03
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</cdnaseq>|
AA = <aaseq>MAAMMASITSELLFFLPFILLALLTFYTTTVAKCHGGHWWRGGT TPAKRKRMNLPPGAAGWPLVGETFGYLRAHPATSVGRFMEQHIARYGKIYRSSLFGER TVVSADAGLNRYILQNEGRLFECSYPRSIGGILGKWSMLVLVGDPHREMRAISLNFLS SVRLRAVLLPEVERHTLLVLRAWPPSSTFSAQHQAKKFTFNLMAKNIMSMDPGEEETE RLRREYITFMKGVVSAPLNLPGTPYWKALKSRAAILGVIERKMEERVEKLSKEDASVE QDDLLGWALKQSNLSKEQILDLLLSLLFAGHETSSMALALAIFFLEGCPKAVQELREE HLGIARRQRLRGECKLSWEDYKEMVFTQCVINETLRLGNVVRFLHRKVIKDVHYKGYD IPSGWKILPVLAAVHLDSSLYEDPQRFNPWRWKSSGSSGGLAQSSSFMPYGGGTRLCA GSELAKLEMAVFLHHLVLNFRWELAEPDQAFVFPFVDFPKGLPIRVHRIAQDDEQE</aaseq>|
DNA = <dnaseqindica>1..266#435..759#863..1015#3829..4080#4229..4321#5114..5192#5290..5396#6284..6529#atggccgccatgatggcgtccataaccagcgagctgctcttctttctccccttcatcctccttgccctgctcacgttctacaccaccaccgtggccaaatgccacggcgggcactggtggcgaggtgggacgacgccggcgaagaggaagcggatgaacctgccgcccggcgccgccgggtggccgctcgtcggcgagacgttcggctacctccgcgcccaccccgccacctccgtcggccgcttcatggagcagcacatcgcacggtcggtggtttcattactcggtacacgcacacagctatataagcacgcgcactgctcgcctcgccgtgaagcaagcttagctagtagctaggatatcgatcgatcggtaataataggctgacggcgagctggggggtggcgtgcggttgcgtgcgtgcgtgcatgcaggtacgggaagatataccggtcgagcctgttcggggagcggacggtggtgtcggcggacgcggggctcaaccggtacatcctgcagaacgaggggaggctgttcgagtgcagctacccgcgcagcatcggcggcatcctgggcaagtggtccatgctggtcctcgtcggggacccgcaccgcgagatgcgcgccatctccctcaacttcctctcctccgtccgcctccgcgccgtcctcctccccgaggtcgagcgccacaccctcctcgtcctccgcgcctggcccccttcctccaccttctccgctcagcaccaagccaagaaggtacaacacacatcgtcctccattaagttgagcatgcatatatcaatcagctcgctctagcgatcaatatcaatgaaacgcgcgtttgcgtgcgtgcgtgcagttcacgttcaacctgatggcgaagaacataatgagcatggacccgggggaggaagagacggagcggctgcggcgggagtacatcaccttcatgaagggcgtggtctccgcgccgctcaacctgcccgggacgccctactggaaggctctcaaggtacgtactcgtagctactagctactccctccgtttcgaaatgtttgacgccgttgactttttatcacatatttgaccgttcgttttatttaaaaaaattaagtaattattaattattttcctatcatttgattcattgttaaatatatttttatgtaggcatataattttacatatctcacaaaagtttttaaataagacgaacggtcaaacatttgctaaaaagtcaacggtgtcaaatatttcgaaacggaggaagtagctagctaccaccccacccgtgcggattgatcgatcgatcgatgtccctctctgcctacacacaccctgcgccccggccgtacgcgcgcaaagtcgtgcgcgcatgagttttactccttccgtttcaaaaaaaaaacgacctagacaagtaatatgatgtaacctagaacaacgaatctatacaggagtatatccaaattcattgtcctagattacatcatatctttgtctatgttgccttttttaggaggagagagtactttacatatgagttatgaccggccggccaggttcctttcttcccctattccgtatgcatactataagcagctgcatataggcatatactactatacccagcttagctaaagcaaacaacttgcaacgacgatcatgtgcaggtccctgttgacagaggttagagatgtaaaccatttttgttgttttcattatctaaagagaaaacaagttactacgacgatcgagatgtacgcaaggggatcatatttatattggagaggagagagagatgatttgcgacttgtgagtgataagtgaatatatatactatatatttgtgtaggagtgacgatgacttccctgtcaatgttttgtttggtatgcacacggggggaaaagagtttagagaaaggaattggctagagagggccaggtttagtttctttaatttttgtttggtggttttgtgtgtagctagtccgcatgcatgatatatcatatatatgcatggcccatcattagatatatatacatgacggcagtgcacttattatatatatgtaggggcatatacaaggtgtatgtgtttagaattgtggatggtccaggaattgttgcagctagttctatactagcatagtgaattagtgatccatatcgagatcaggtccaatcggtcagctcgcacagcgggtcttttagtacgatttgttcggacaaattatcttgcctttgagataatcaacataggattgaaactgcatactagcaatagaccatagctatttggcaatatagaaagctaaaaaatgttgattgaaaggtgcgtgcatgcatgctagcagtatgtcttttaggccaggtccagtttttttttttttctcaaaggcagttgcttaaggtgaaattctagcagaggtcatatgaatatatttattatcttccaaacatgagccaaaacctatatatagttaactcaaaaaaccatgcattggtacttcttctgagtatgcatcctggggactgatcgatcatgcagattcttttctttctttttccctggggacgtactgatcatcactcatgtgagcataaaatgttgcctactttttactgcatgtgaggtcacactcaaatgatctttattgattgcctcagcatgcatgctgctagctttcacgatctctctccgatacacacaccctgtgctgtgctcaacagcatgcaacctggcaggtggaatttttaatctgttggaaagtaaatgaccaaccttcataatttggcatgattatatgctggggaatctatcatttatcatgattcaaccgctgaccaaactctttgccacacacccgaaccacattaatttcgttaagcggatgattagccagcaaccttcacagtttggttttaattagttaatgaaagggtgcttgtagttttattagctgaagagacactgcagtactcctactgagcgttctccttaattgggaaatagatctctgcagtactatcctttaactttaagtgaataacatgtagctaggtctcacacttatcaggcccacacatcagtgatccaagtcaaatggcaatacaccataggagattcttccactaattggctaacaaaaataaactgtgatcactgatcacagctgctaacactataacacattaatttgcatgtctcatgaaatgccagatgagttggattaattatctatcttatgttcagtgccgaggaatatacaaggaaacatcattggtatagttgtggagttctgagtgcattatctcacccagtacggtcaaattaatcctagtctccgtaaatatggttgtgtgcactcttagttaattagtgtagacccggaaattatactttgtcgagaatgttgttatccgtgtgttaagtttcaagtaccctgcaggcctataactaaacactgccaacatggatgtttaagaatgaattatacatacagcgcacttggaaaagaaaaggggtagctacccagctgaagaacacctctctctcatttgaaggtcgacacaatatgtgtcttttctatatcgaaaggaaacgacattttttctcttcaaaacagaaagctatgacaagagacataatcctagactagtaggcggttaggcaacaccatatctgaatttgataggtgtcccacatattttttttcttttcctgataagaaggaacttttcaccgtgagcttatttttttctgttctatgtcgacattagttccatgatgtttactgtgtatttttactgttcccctgcagtcgcgtgctgccattctcggagtaatagagaggaaaatggaagagcgggttgagaagctgagcaaggaggatgcaagcgtagagcaagacgatcttctcggatgggctctgaaacaatctaacctttcaaaagagcaaatcctggacctcttgctgagcttgctcttcgccgggcacgagacgtcgtccatggcgctcgccctcgccatcttcttccttgaaggctgccccaaggctgtccaagaactgagggtacagagaattacagattcgaaattaacatctactgtcactagctcagtgctacgtttaactgtagagcaacaattgcagaaagcgaaggcatcagaaatgacatgggaatcgattagttacactcacactgacgcgtgacatgcaggaggagcatcttgggattgcaaggagacaaaggctaagaggggagtgcaaattgagctgggaagactacaaagagatggttttcacgcaatgtgtaaggccccccctactttgttccttccgatccctcctccctaaacgctactaatataaatcggtgcaaaggctttgtgctagcgtagctcttgctgttttgtttggtcacttgcgtagagagatagcaacttcccgaaatgtcatgtacgaatggagactgaaagctctgcgaaataattgccgggccacgtactacagtacttacgctccctctcttcagtttggatcgatcgagctaagaaggcgcacgcatgtgtgcattgatcgatagagtcgatctgttctgccttgtaagctagccagctgaggttctttcgccttggaagtttctacagggtgaggcccatcgtcttgcaacgatccaaatgccatgacaagtacctgttgttgtaggcttccagtagcacggcctgcccccaagaaggtatctgaatcttctgcattgctgttcagacggcagagcagaacatgccgaaataccattccagtcagttgcagtatatgtgtgtggtggtatcagataaaggagcagagtaatttgaggccttagaaattcagatgagttgctcgacaattgtcgcctaccgatcgagaataagttgcatgcatgtatgtgatgtaattatgggtcttagtcagcaggatagtttagtcgctgcagtactgtagattgtagtacttgcgttcaaagcagacccaaaacagaacttgcattcaggaaacaaaacatgaaaaaagaaatatacagtacttggcactgatatttcttttgtttgggtccttattgcaggtcataaacgagacgttgcggctaggaaacgtggtcaggttcctgcaccggaaggtcatcaaggacgtgcactacaagggtgagtagctcgatttcactcttcttccctgaatcgacctctcctctatctgacagcagaagctaactgaactgaactgaactgaactgaactgaaggttatgacattccaagcggatggaagatcctgccggtgttagccgcggtgcatctggactcgtccctgtacgaggacccccagcgcttcaatccctggagatggaaggtcagtcgcagtaggattatcagtgaatctaatctattaatctcggtttaattgctgggattcaaggaaaagaaaatactgggacaagcgcggatgtcgctgttgggcagtgtctgtcaatcataagcgaaggggatcatgtcagaatctgcaaaagcagcacaggctaaacgcgtgcttgcctcgagcagcaatcatgcagagaggcaagctccgcgataggaacatgcatggtgaatcactgaacctgatatgaaagtgaatgctcagctcagctcgctcgctcgcgcgcgcatgtgctttctcatttcgccgtgttgcgtgctagctagtcaatggtgttcccacgaattgttttctcttctcatcacacaaagtcagtggtgaaaaccgaggtgcatgtgcggtaatggaacgggggattaggcggcgacgccgtgatggattatgcattcgtgcgagaaaaatccaggagcgttggtccaaatcgcaagcggatcaaaccgaaccaatcagtggacgattggagccctgcagtccatccatcacatctccattcttatccgctgcgactcacggacattagttgttgaggggtgaacgacgcatgtgtgtatgtatgctcgatcgatcgatcgaacaatggatctgtgtgtggtgtagttgccaacgccgacgagattatctgctcgtctgcatgcagccgcgaaaaagacatgtgtccggattatcctgcaggagattgagcaagataatctgctgcgcaaaaactgcagccaatcgatcattccatgacgcgaaaaaaaaaacaataattactactgctgatacgataacttcgcttcgaattgttgacacacaagtgttcttcttttgttcgattgcagagtagcggatcatccggcggcttggctcagagcagcagcttcatgccgtacggcggcgggacgcggctgtgcgccgggtcggagctcgcgaagctggagatggccgtgttcttgcaccacctggtgctcaacttcaggtgggagctcgccgagccggaccaagccttcgtcttccccttcgtcgacttccccaagggccttcccattagggttcatagaattgcacaggatgatgagcaggagtaa</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001055982.2 RefSeq:Os03g0227700]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 3]]
[[Category:Chromosome 3]]

Os03g0227700

2014-05-24T11:11:47Z

LQ: /* Annotated Information */

Os01g0197100

2014-05-24T10:55:20Z

LQ: /* Annotated Information */

Please input one-sentence summary here.

==Annotated Information==
This gene is a rice dwarf mutant, ebisu dwarf (d2), which first was described asebisu dwarf in an article published in 1925. The D2 gene encodes a P450 protein that is classified in the CYP90D group that is highly similar to other BR biosynthesis P450 proteins, such as CPD/CYP90A, DWF4/CYP90B, and DWARF/CYP85. ebisu dwarf (dwarf2 or d2) is a good example of dwarf mutant, although its dwarfism is slightly stronger than the desirable level. In fact, the erect leaves of d2 allow this cultivar to be planted more densely than the original cultivar, which has bent leaves; consequently, a greater volume of crop products can be harvested in the same cultivation area.This dwarf mutant has unusual phenotypic characteristics, such as its erect leaves and the specific inhibition of second internode elongation. Thus, elucidation of the molecular mechanism of the relationship between dwarfism and erect leaves in d2 mutants is important for further molecular breeding for architectural modification.

===Expression===
Please input expression information here.

===Evolution===
Please input evolution information here.

You can also add sub-section(s) at will.
We characterized a rice dwarf mutant, ebisu dwarf (d2). It showed the pleiotropic abnormal phenotype similar to that of the rice brassinosteroid (BR)-insensitive mutant, d61. The dwarf phenotype of d2 was rescued by exogenous brassinolide treatment. The accumulation profile of BR intermediates in the d2 mutants confirmed that these plants are deficient in late BR biosynthesis. We cloned the D2 gene by map-based cloning. The D2 gene encoded a novel cytochrome P450 classified in CYP90D that is highly similar to the reported BR synthesis enzymes. Introduction of the wild D2 gene into d2-1 rescued the abnormal phenotype of the mutants. In feeding experiments, 3-dehydro-6-deoxoteasterone, 3-dehydroteasterone, and brassinolide effectively caused the lamina joints of the d2 plants to bend, whereas more upstream compounds did not cause bending. Based on these results, we conclude that D2/CYP90D2 catalyzes the steps from 6-deoxoteasterone to 3-dehydro-6-deoxoteasterone and from teasterone to 3-dehydroteasterone in the late BR biosynthesis pathway[1].

==Labs working on this gene==
Please input related labs here.

==References==

1. Zhi Hong;Miyako Ueguchi-Tanaka;Kazuto Umemura;Sakurako Uozu;Shozo Fujioka;Suguru Takatsuto;Shigeo Yoshida;Motoyuki Ashikari;Hidemi Kitano and Makoto Matsuok
A Rice Brassinosteroid-Deficient Mutant, ebisu dwarf (d2), Is Caused by a Loss of Function of a New Member of Cytochrome P450
The Plant Cell, 2003, 15(12): 2900-2910

==Structured Information==
{{JaponicaGene|
GeneName = Os01g0197100|
Description = Similar to Cytochrome P450 90C1 (EC 1.14.-.-) (ROTUNDIFOLIA3)|
Version = NM_001048832.1 GI:115435077 GeneID:4327329|
Length = 7389 bp|
Definition = Oryza sativa Japonica Group Os01g0197100, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 1|Chromosome 1]]|
AP = Chromosome 1:5235623..5243011|
CDS = 5235623..5235873,5235958..5236285,5236786..5236935,5237255..5237503,5238581..5238670 ,5238781..5238966,5239477..5239598,5242915..5243011|
GCID = <gbrowseImage1>
name=NC_008394:5235623..5243011
source=RiceChromosome01
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008394:5235623..5243011
source=RiceChromosome01
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggtgtcggcggccgccggttgggcggcgccggcgtttgcggtcgccgccgtggttatttgggtggtgctgtgtagtgagctcctgcggaggcggcggcgtggtgcaggcagcggcaagggggacgcggcggcggcggcgcgcctcccgccggggagcttcgggtggccagtggtgggcgagacgctggagttcgtgtcgtgcgcctactcgccgcgccccgaggcgttcgtcgacaagcgccggaagctgcacgggagcgcggtgttcaggtcgcacctgttcgggtcggcgacggtggtgacggcggacgcggaggtgagccggttcgtgctgcagagcgacgcgcgggcgttcgtgccgtggtacccgcggtcgctgacggagctgatgggcaagtcctccatcctcctcatcaacggcgcgctccagcgacgcgtccacggcctcgtcggcgccttcttcaagtcctcccacctcaagtcccagctcaccgccgacatgcgccgccgcctctcccccgccctctcctccttccccgactcctccctcctccacgtccagcacctcgccaagtcggtggtgttcgaaatcctggtgaggggcctgatcgggctggaggcaggggaggagatgcagcagctgaagcagcaattccaggaatttattgtcggcctcatgtccctccccattaagctgcctggcactaggctctacagatcactccaggccaagaagaagatggcgaggctgatacagaggatcatccgggagaagagggcaaggagggccgccgcctcgccgccgcgggacgccatcgacgtgctcatcggagacggcagcgatgagctcaccgacgagctcatctccgacaacatgatcgacctcatgatccccgccgaggactccgtcccggtgctcatcacgctcgccgtcaagttcctcagcgagtgccctctcgccctgcaccaactggaagaggagaacatacagctcaagaggcgaaaaaccgacatgggtgagaccttgcaatggacagactacatgtcattgtcattcacacaacatgtgataacagagacgctgcggttgggcaacatcatcggtgggatcatgcgcaaggcggtgcgcgacgtcgaggtgaaggggcacctcatccccaaggggtggtgcgtgtttgtgtacttccggtcagtccacctcgatgatacgctctacgatgagccctacaagttcaacccatggaggtggaaggagaaggacatgagcaatggcagcttcactccttttggtggtgggcagaggctgtgcccaggcctggatctggccaggctggaagcttccatcttccttcaccacttggtcaccagcttcaggtgggtggcggaggaggaccacatcgtcaacttccccaccgtgcggctcaagcggggcatgcccatcagggtcaccgccaaggaggacgacgactag</cdnaseq>|
AA = <aaseq>MVSAAAGWAAPAFAVAAVVIWVVLCSELLRRRRRGAGSGKGDAA AAARLPPGSFGWPVVGETLEFVSCAYSPRPEAFVDKRRKLHGSAVFRSHLFGSATVVT ADAEVSRFVLQSDARAFVPWYPRSLTELMGKSSILLINGALQRRVHGLVGAFFKSSHL KSQLTADMRRRLSPALSSFPDSSLLHVQHLAKSVVFEILVRGLIGLEAGEEMQQLKQQ FQEFIVGLMSLPIKLPGTRLYRSLQAKKKMARLIQRIIREKRARRAAASPPRDAIDVL IGDGSDELTDELISDNMIDLMIPAEDSVPVLITLAVKFLSECPLALHQLEEENIQLKR RKTDMGETLQWTDYMSLSFTQHVITETLRLGNIIGGIMRKAVRDVEVKGHLIPKGWCV FVYFRSVHLDDTLYDEPYKFNPWRWKEKDMSNGSFTPFGGGQRLCPGLDLARLEASIF LHHLVTSFRWVAEEDHIVNFPTVRLKRGMPIRVTAKEDDD</aaseq>|
DNA = <dnaseqindica>1..251#336..663#1164..1313#1633..1881#2959..3048#3159..3344#3855..3976#7293..7389#atggtgtcggcggccgccggttgggcggcgccggcgtttgcggtcgccgccgtggttatttgggtggtgctgtgtagtgagctcctgcggaggcggcggcgtggtgcaggcagcggcaagggggacgcggcggcggcggcgcgcctcccgccggggagcttcgggtggccagtggtgggcgagacgctggagttcgtgtcgtgcgcctactcgccgcgccccgaggcgttcgtcgacaagcgccggaagctgtaagcctagccacctccgccgccgccgccgtcccgaggcgttcggctctgactgacgggcatgggtgtgcatgcatggtgcaggcacgggagcgcggtgttcaggtcgcacctgttcgggtcggcgacggtggtgacggcggacgcggaggtgagccggttcgtgctgcagagcgacgcgcgggcgttcgtgccgtggtacccgcggtcgctgacggagctgatgggcaagtcctccatcctcctcatcaacggcgcgctccagcgacgcgtccacggcctcgtcggcgccttcttcaagtcctcccacctcaagtcccagctcaccgccgacatgcgccgccgcctctcccccgccctctcctccttccccgactcctccctcctccacgtccagcacctcgccaagtcggtacgtcctccttcttctccctctccggcgagctcctcgacgagatagagtgggtggatgaattggaggaggagcagagtgggtgggcgtgggcgagcgccggcgtgcgtgcacacgtgcgcgcccgtgtcaccacatggaagtaacttacacaaggccgggaaagggaaggcccaaaaggagtgggcccaatcacacacactctctctctctctctctctctctctctctcatctcatattctgactctagagagagagagagtggctcacctccatgtgggccccaccccatcggagtagagttaccactaccagcagccgaaaatccacgttcctgttcgccaccagagcgtgtgtttgtgcactcgtaaatgtttattttttctctgcgtattcctctggaagagatgcaaatacagtcttgaacatttgtaatttttagagatgaggatggaacgaaatgtaattgcaagaaatggtgatgaaatgcaaatgcaggtggtgttcgaaatcctggtgaggggcctgatcgggctggaggcaggggaggagatgcagcagctgaagcagcaattccaggaatttattgtcggcctcatgtccctccccattaagctgcctggcactaggctctacagatcactccaggtactctctctctctctctctccctctcatctaagttctctttctctctctacctgtagtactccatgtaaacctgcactgcaccacacattgatctctactactctcttctgtcagccactctgtctctcctttatgtctgatgttgctcagccctactccatgcagcatcagcagcagcatgatcagtgtctcagctgccatcaccattaacctctcttaatatgttctctctgtttcacaatcttggattaaatgtatacaattgcactcgatccattgcacatttttgacacattattgtgtgtgtggtcatcaggccaagaagaagatggcgaggctgatacagaggatcatccgggagaagagggcaaggagggccgccgcctcgccgccgcgggacgccatcgacgtgctcatcggagacggcagcgatgagctcaccgacgagctcatctccgacaacatgatcgacctcatgatccccgccgaggactccgtcccggtgctcatcacgctcgccgtcaagttcctcagcgagtgccctctcgccctgcaccaactggaagtaagcctgtatacctgaaacctctcttacaacacagtaagctcttaattggatcaatacctgcatataattagatgctagagatgaatcctgaacagaaaagaaaaataaaaccataccctaggaagtaattaaggatcctctgttgctgggcatgggcagtcagcaaaaagggtccagaactgaaaagagttggattttctttcttttttgtctgaaaggaacatgatcaactattcattttgtcagagccatgtgttgtgcatgatatactactatcactattgtctactacacacaatggcagaggatctgtgtccccaaacaaaaactagatcataaaaagcctacgaataagggagagatacgtaggtgggcagaagggatcagcagcatataaagggttgtgccaaccttggtccatccctttcttgttggcatcttagagctatcattattatttgttatctaatgtgccatctgaaaagaaatactactactacttgtgtttggaccaatctttgcaagatgacgtgcgttgcatcattgtcaatgtcaggtcgggtacatacatatcattagaacagagtcgccttctcacaaaagaaaaggcttactatcatgatgtatcatctacttacatatcattagtttgtgctattatgcgttttttttttctaaccggctataaacgcatgtgcaaggaatagtttttttggttgaagtatggagggttttttatcaactggggaaaaaagaaggttacataatttggctatttattctatggagattagaaggcagcagtaatatttcctagctagcatgtcctattacaccagtagcattgtgtcatatgctagattccatttacatcactacagtgctaagaaattactcgcgtgtcgactgtgtttctgttttgatgccatgtgagcaataaactcattgaaacctctgcatgtctaccactgttagagaacatgcatatcgcggctgaagataaaatccaccgtcagtaattacttgttgtaacaatgatgaataataacaccaatcacattctaatgctaccaggaggagaacatacagctcaagaggcgaaaaaccgacatgggtgagaccttgcaatggacagactacatgtcattgtcattcacacaacatgtaagtatagttcattaacactgtaacataaatttctatagctcaaaatctactggccccgtgtggtttttttcccatgaccccatgcgatttggaaaattggagtgtaggtgataacagagacgctgcggttgggcaacatcatcggtgggatcatgcgcaaggcggtgcgcgacgtcgaggtgaaggggcacctcatccccaaggggtggtgcgtgtttgtgtacttccggtcagtccacctcgatgatacgctctacgatgagccctacaagttcaacccatggaggtggaaggtaagaaagagccccacatttggtagagatgttcatccaattgctacccctttaggcaacaccagcataataaggatttgatgagtgcggttttggtgggactcctaacatgtgggcttacttctatatcagtaatctcatgatggatttgacattgccttgtactaacacttagtattgcagaaaataacagagattccctcaaagttgcattagtgattgatatatacacctaggcagaagtcaattgaatctcccaaaataactggatttggtagttctttgaactggtaaattcagacaacataagcttaagctgtccaggaaaggaatacacaccctttttttccctctattgtgtactttgtgtaagctctgaaaatgatatgcaaataattaatctaggcaaatgatggtgctgtactactgttgaggcatcctatcattatgatttgatttgggaagtctaataatgtgctaattggtcaagaaatcaatcaatctcaacaggagaaggacatgagcaatggcagcttcactccttttggtggtgggcagaggctgtgcccaggcctggatctggccaggctggaagcttccatcttccttcaccacttggtcaccagcttcaggtatggatcaccacatctcaatcttggccattcttagtgcagccattgattccaactccttagctttgtttcatgatcacttggcaacaatagcttttttttctttctagataatggaatacaaactacaacttttgcataattttccatgatcgctcggcaccgatagcatttttttctagataaaagaataaaaatcacgacctctacatcatataatgaatggaataaaaatcacggcctttgcatcatataatgcacgcaaccgggcaccaatggcatgcatctcatgtagtcatgtccctatcatttggccatgccctcttcgcatgcacattcatgttcctagtaataactaatctctgtagaacaatagaagagcggaacagtagatgaattatgcgttgattgacagaataattgtgggggggtatatagatcgggtacgtacatgaggatttgaagatcaagccgaatctttccagagataaaaaaaaaagaaaaaaattatgtcagataataactggtcctaatacgtcactagggatagcatcacgaggctttctcccatgcttatacctcctctgttatcatcttccttgagggagtgatcttactgtgcttaaggaggtatctaaagcggtggtgaagccagcgacatcgtcgagcccggataactaatagtgttttgtagtgatcaatccatttttacacgcgtcaacggtgattggtagtggagacgccgacagcgaagcctagacggtcgttcgacctggagccgtctccacctctgatcgaaagatatagtatggtattttcgcctctcacactgggagatacaactttcggcctacaaaatcaacttctggttttaaatctaagatatagattatttctagattcaagcttagaaattgttttttttaagagacagagatagtaccaaattctatttctataattttcttaaatacagtaaattatttcttctcgcgaggaggattacactgcgtaggaacactaggtcatttttcatgtgaaagaaaaaaatggaagctacgaggaagacaaataatttcggcgcgcggggcgaaaaagaagaagatcaaggaagatacccaaggtgggaggtcggtccagcctctattgcgaatagagcggggtgttgttccgcgcgtggggccacacgtgcgcgcacatgcggcgcatatccatcccttcccccccacacccacatgtgtgcatgcatgcggactacgccgctgtacgcgtgcgtggcttacggcgcccgctgctacagctacagcgcccgcacgtgtcagctaccacgccacgccctgcatgcccaccgccgctgcacctccccttgagacgtggtcgctgggcggtggtgggtctcacatgtcagtacgtctttcgtcaaaaaaaaaattcaactttaatttgtttttttcatggtctcacaaccatgtagcttaccaatgcaaacagactaatctaaaaaaaaaaaaataagcttagctataatgcacgcttcgattaagcaatgttacgatgcgtgtgcttagctcaggatagtgcctatattgacacgaaaatatactacatccatataaaaaaatataagcattttttactataaatttatatatttattcatatttatagctaaaaatacttatattttgggtcggagcgagtatcatattttgattaccagtcgtccatcagctatggagtctgtatagggctacgtcctacgttcgggaggtagaaaatatggtgatttattttatttttttgcggggaaatatggtgatttattaatatatgattaattaagtattagttaaaaacttaaagaataggttaatataaatttaaaaataatttttatataaaaaatcgtattgtttagtagtttagaaaatgtgccggatcgggttgagttgggaaaggaggcgggaagaagcgtggtgtagtaattggcgaagggggtaagaagggggagaccggagacatgggtgtgatggttaggcagccggaatgggctctctatcgactcgaacagttggatgagatgagacgagatgggtgacatgcgtgcgtgcatgcatactttgtgccttgtccggtccgtgactgaactgaatcccaggaacgcgaatataatactgtagcacgcacgcacgcacgcacgcatccagctgatctagccggccgagatgtgccactttgaccggtcgtcagacacgctgcatcaatgtaggagcacgtgccagggtatggtcgctgcatgcctgcctgcaaacaatactatactcctcccataaacaaatcaaactccaacaagagaatccgacttgctagaaaaacatcatatcactttaaacaatggagattggagagtccatcagtccaatgtgtttttctgtcttctttgaccatcttccaattccccattatgcttgtgctgacctctatctaattgatcctaagagaagcgttttgcaacatcttgtaaaaaagcaaatcacataatattactcgtgctttatgtagctaagctacattggatcaggatcctctgcagtcaaagtcacgtgactttgtcactgacattgtgggtccgataatcattgggtccacatgtcttacaaagacaatccggctacatcatcatgccagtttccgtaataaattgaccagtaatttagtactatggactttagaacctggcacatgcatagcacagattcatgcatgcgtgcactgcatctgcacgcatttgcattggttgcacaagcagctagctggcctacatataacatattactgtgttatgcgagcaactggccactgtccacatgaagccagtaaatcaactatcaacattgatagtagtaaagctcttgttcagagttgcaggcatataggtgctccctgcatggtttgctcaaacttaagatagatgatactacattctaaaaatgcaagtatttctggtatgtatctggacaagatggttaggactctagctataacatagaaaatgcttatattagaatacacttcattcgtctcaaaatataaggtattttgatcggatatcgatcccatccggttaaataccttaagtagggagtggcagtggcatcaccatcgatcatgtgttattgataaatgattatgattatgagatctcttatctacaatctctttctgaataatcgacctgctgatccatccattgcatgcatggttgttgcctgacaaggacattgaaatggactgctaattgctgccatggatgcaggtgggtggcggaggaggaccacatcgtcaacttccccaccgtgcggctcaagcggggcatgcccatcagggtcaccgccaaggaggacgacgactag</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001048832.1 RefSeq:Os01g0197100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 1]]
[[Category:Chromosome 1]]

Os01g0197100

2014-05-24T10:54:22Z

LQ: /* Annotated Information */

Please input one-sentence summary here.

==Annotated Information==
This gene is a rice dwarf mutant, ebisu dwarf (d2), which first was described asebisu dwarf in an article published in 1925. The D2 gene encodes a P450 protein that is classified in the CYP90D group that is highly similar to other BR biosynthesis P450 proteins, such as CPD/CYP90A, DWF4/CYP90B, and DWARF/CYP85. ebisu dwarf (dwarf2 or d2) is a good example of dwarf mutant, although its dwarfism is slightly stronger than the desirable level. In fact, the erect leaves of d2 allow this cultivar to be planted more densely than the original cultivar, which has bent leaves; consequently, a greater volume of crop products can be harvested in the same cultivation area.This dwarf mutant has unusual phenotypic characteristics, such as its erect leaves and the specific inhibition of second internode elongation. Thus, elucidation of the molecular mechanism of the relationship between dwarfism and erect leaves in d2 mutants is important for further molecular breeding for architectural modification.

===Expression===
Please input expression information here.

===Evolution===
Please input evolution information here.

You can also add sub-section(s) at will.
We characterized a rice dwarf mutant, ebisu dwarf (d2). It showed the pleiotropic abnormal phenotype similar to that of the rice brassinosteroid (BR)-insensitive mutant, d61. The dwarf phenotype of d2 was rescued by exogenous brassinolide treatment. The accumulation profile of BR intermediates in the d2 mutants confirmed that these plants are deficient in late BR biosynthesis. We cloned the D2 gene by map-based cloning. The D2 gene encoded a novel cytochrome P450 classified in CYP90D that is highly similar to the reported BR synthesis enzymes. Introduction of the wild D2 gene into d2-1 rescued the abnormal phenotype of the mutants. In feeding experiments, 3-dehydro-6-deoxoteasterone, 3-dehydroteasterone, and brassinolide effectively caused the lamina joints of the d2 plants to bend, whereas more upstream compounds did not cause bending. Based on these results, we conclude that D2/CYP90D2 catalyzes the steps from 6-deoxoteasterone to 3-dehydro-6-deoxoteasterone and from teasterone to 3-dehydroteasterone in the late BR biosynthesis pathway.

==Labs working on this gene==
Please input related labs here.

==References==

1. Zhi Hong;Miyako Ueguchi-Tanaka;Kazuto Umemura;Sakurako Uozu;Shozo Fujioka;Suguru Takatsuto;Shigeo Yoshida;Motoyuki Ashikari;Hidemi Kitano and Makoto Matsuok
A Rice Brassinosteroid-Deficient Mutant, ebisu dwarf (d2), Is Caused by a Loss of Function of a New Member of Cytochrome P450
The Plant Cell, 2003, 15(12): 2900-2910

==Structured Information==
{{JaponicaGene|
GeneName = Os01g0197100|
Description = Similar to Cytochrome P450 90C1 (EC 1.14.-.-) (ROTUNDIFOLIA3)|
Version = NM_001048832.1 GI:115435077 GeneID:4327329|
Length = 7389 bp|
Definition = Oryza sativa Japonica Group Os01g0197100, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 1|Chromosome 1]]|
AP = Chromosome 1:5235623..5243011|
CDS = 5235623..5235873,5235958..5236285,5236786..5236935,5237255..5237503,5238581..5238670 ,5238781..5238966,5239477..5239598,5242915..5243011|
GCID = <gbrowseImage1>
name=NC_008394:5235623..5243011
source=RiceChromosome01
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008394:5235623..5243011
source=RiceChromosome01
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atggtgtcggcggccgccggttgggcggcgccggcgtttgcggtcgccgccgtggttatttgggtggtgctgtgtagtgagctcctgcggaggcggcggcgtggtgcaggcagcggcaagggggacgcggcggcggcggcgcgcctcccgccggggagcttcgggtggccagtggtgggcgagacgctggagttcgtgtcgtgcgcctactcgccgcgccccgaggcgttcgtcgacaagcgccggaagctgcacgggagcgcggtgttcaggtcgcacctgttcgggtcggcgacggtggtgacggcggacgcggaggtgagccggttcgtgctgcagagcgacgcgcgggcgttcgtgccgtggtacccgcggtcgctgacggagctgatgggcaagtcctccatcctcctcatcaacggcgcgctccagcgacgcgtccacggcctcgtcggcgccttcttcaagtcctcccacctcaagtcccagctcaccgccgacatgcgccgccgcctctcccccgccctctcctccttccccgactcctccctcctccacgtccagcacctcgccaagtcggtggtgttcgaaatcctggtgaggggcctgatcgggctggaggcaggggaggagatgcagcagctgaagcagcaattccaggaatttattgtcggcctcatgtccctccccattaagctgcctggcactaggctctacagatcactccaggccaagaagaagatggcgaggctgatacagaggatcatccgggagaagagggcaaggagggccgccgcctcgccgccgcgggacgccatcgacgtgctcatcggagacggcagcgatgagctcaccgacgagctcatctccgacaacatgatcgacctcatgatccccgccgaggactccgtcccggtgctcatcacgctcgccgtcaagttcctcagcgagtgccctctcgccctgcaccaactggaagaggagaacatacagctcaagaggcgaaaaaccgacatgggtgagaccttgcaatggacagactacatgtcattgtcattcacacaacatgtgataacagagacgctgcggttgggcaacatcatcggtgggatcatgcgcaaggcggtgcgcgacgtcgaggtgaaggggcacctcatccccaaggggtggtgcgtgtttgtgtacttccggtcagtccacctcgatgatacgctctacgatgagccctacaagttcaacccatggaggtggaaggagaaggacatgagcaatggcagcttcactccttttggtggtgggcagaggctgtgcccaggcctggatctggccaggctggaagcttccatcttccttcaccacttggtcaccagcttcaggtgggtggcggaggaggaccacatcgtcaacttccccaccgtgcggctcaagcggggcatgcccatcagggtcaccgccaaggaggacgacgactag</cdnaseq>|
AA = <aaseq>MVSAAAGWAAPAFAVAAVVIWVVLCSELLRRRRRGAGSGKGDAA AAARLPPGSFGWPVVGETLEFVSCAYSPRPEAFVDKRRKLHGSAVFRSHLFGSATVVT ADAEVSRFVLQSDARAFVPWYPRSLTELMGKSSILLINGALQRRVHGLVGAFFKSSHL KSQLTADMRRRLSPALSSFPDSSLLHVQHLAKSVVFEILVRGLIGLEAGEEMQQLKQQ FQEFIVGLMSLPIKLPGTRLYRSLQAKKKMARLIQRIIREKRARRAAASPPRDAIDVL IGDGSDELTDELISDNMIDLMIPAEDSVPVLITLAVKFLSECPLALHQLEEENIQLKR RKTDMGETLQWTDYMSLSFTQHVITETLRLGNIIGGIMRKAVRDVEVKGHLIPKGWCV FVYFRSVHLDDTLYDEPYKFNPWRWKEKDMSNGSFTPFGGGQRLCPGLDLARLEASIF LHHLVTSFRWVAEEDHIVNFPTVRLKRGMPIRVTAKEDDD</aaseq>|
DNA = <dnaseqindica>1..251#336..663#1164..1313#1633..1881#2959..3048#3159..3344#3855..3976#7293..7389#atggtgtcggcggccgccggttgggcggcgccggcgtttgcggtcgccgccgtggttatttgggtggtgctgtgtagtgagctcctgcggaggcggcggcgtggtgcaggcagcggcaagggggacgcggcggcggcggcgcgcctcccgccggggagcttcgggtggccagtggtgggcgagacgctggagttcgtgtcgtgcgcctactcgccgcgccccgaggcgttcgtcgacaagcgccggaagctgtaagcctagccacctccgccgccgccgccgtcccgaggcgttcggctctgactgacgggcatgggtgtgcatgcatggtgcaggcacgggagcgcggtgttcaggtcgcacctgttcgggtcggcgacggtggtgacggcggacgcggaggtgagccggttcgtgctgcagagcgacgcgcgggcgttcgtgccgtggtacccgcggtcgctgacggagctgatgggcaagtcctccatcctcctcatcaacggcgcgctccagcgacgcgtccacggcctcgtcggcgccttcttcaagtcctcccacctcaagtcccagctcaccgccgacatgcgccgccgcctctcccccgccctctcctccttccccgactcctccctcctccacgtccagcacctcgccaagtcggtacgtcctccttcttctccctctccggcgagctcctcgacgagatagagtgggtggatgaattggaggaggagcagagtgggtgggcgtgggcgagcgccggcgtgcgtgcacacgtgcgcgcccgtgtcaccacatggaagtaacttacacaaggccgggaaagggaaggcccaaaaggagtgggcccaatcacacacactctctctctctctctctctctctctctctcatctcatattctgactctagagagagagagagtggctcacctccatgtgggccccaccccatcggagtagagttaccactaccagcagccgaaaatccacgttcctgttcgccaccagagcgtgtgtttgtgcactcgtaaatgtttattttttctctgcgtattcctctggaagagatgcaaatacagtcttgaacatttgtaatttttagagatgaggatggaacgaaatgtaattgcaagaaatggtgatgaaatgcaaatgcaggtggtgttcgaaatcctggtgaggggcctgatcgggctggaggcaggggaggagatgcagcagctgaagcagcaattccaggaatttattgtcggcctcatgtccctccccattaagctgcctggcactaggctctacagatcactccaggtactctctctctctctctctccctctcatctaagttctctttctctctctacctgtagtactccatgtaaacctgcactgcaccacacattgatctctactactctcttctgtcagccactctgtctctcctttatgtctgatgttgctcagccctactccatgcagcatcagcagcagcatgatcagtgtctcagctgccatcaccattaacctctcttaatatgttctctctgtttcacaatcttggattaaatgtatacaattgcactcgatccattgcacatttttgacacattattgtgtgtgtggtcatcaggccaagaagaagatggcgaggctgatacagaggatcatccgggagaagagggcaaggagggccgccgcctcgccgccgcgggacgccatcgacgtgctcatcggagacggcagcgatgagctcaccgacgagctcatctccgacaacatgatcgacctcatgatccccgccgaggactccgtcccggtgctcatcacgctcgccgtcaagttcctcagcgagtgccctctcgccctgcaccaactggaagtaagcctgtatacctgaaacctctcttacaacacagtaagctcttaattggatcaatacctgcatataattagatgctagagatgaatcctgaacagaaaagaaaaataaaaccataccctaggaagtaattaaggatcctctgttgctgggcatgggcagtcagcaaaaagggtccagaactgaaaagagttggattttctttcttttttgtctgaaaggaacatgatcaactattcattttgtcagagccatgtgttgtgcatgatatactactatcactattgtctactacacacaatggcagaggatctgtgtccccaaacaaaaactagatcataaaaagcctacgaataagggagagatacgtaggtgggcagaagggatcagcagcatataaagggttgtgccaaccttggtccatccctttcttgttggcatcttagagctatcattattatttgttatctaatgtgccatctgaaaagaaatactactactacttgtgtttggaccaatctttgcaagatgacgtgcgttgcatcattgtcaatgtcaggtcgggtacatacatatcattagaacagagtcgccttctcacaaaagaaaaggcttactatcatgatgtatcatctacttacatatcattagtttgtgctattatgcgttttttttttctaaccggctataaacgcatgtgcaaggaatagtttttttggttgaagtatggagggttttttatcaactggggaaaaaagaaggttacataatttggctatttattctatggagattagaaggcagcagtaatatttcctagctagcatgtcctattacaccagtagcattgtgtcatatgctagattccatttacatcactacagtgctaagaaattactcgcgtgtcgactgtgtttctgttttgatgccatgtgagcaataaactcattgaaacctctgcatgtctaccactgttagagaacatgcatatcgcggctgaagataaaatccaccgtcagtaattacttgttgtaacaatgatgaataataacaccaatcacattctaatgctaccaggaggagaacatacagctcaagaggcgaaaaaccgacatgggtgagaccttgcaatggacagactacatgtcattgtcattcacacaacatgtaagtatagttcattaacactgtaacataaatttctatagctcaaaatctactggccccgtgtggtttttttcccatgaccccatgcgatttggaaaattggagtgtaggtgataacagagacgctgcggttgggcaacatcatcggtgggatcatgcgcaaggcggtgcgcgacgtcgaggtgaaggggcacctcatccccaaggggtggtgcgtgtttgtgtacttccggtcagtccacctcgatgatacgctctacgatgagccctacaagttcaacccatggaggtggaaggtaagaaagagccccacatttggtagagatgttcatccaattgctacccctttaggcaacaccagcataataaggatttgatgagtgcggttttggtgggactcctaacatgtgggcttacttctatatcagtaatctcatgatggatttgacattgccttgtactaacacttagtattgcagaaaataacagagattccctcaaagttgcattagtgattgatatatacacctaggcagaagtcaattgaatctcccaaaataactggatttggtagttctttgaactggtaaattcagacaacataagcttaagctgtccaggaaaggaatacacaccctttttttccctctattgtgtactttgtgtaagctctgaaaatgatatgcaaataattaatctaggcaaatgatggtgctgtactactgttgaggcatcctatcattatgatttgatttgggaagtctaataatgtgctaattggtcaagaaatcaatcaatctcaacaggagaaggacatgagcaatggcagcttcactccttttggtggtgggcagaggctgtgcccaggcctggatctggccaggctggaagcttccatcttccttcaccacttggtcaccagcttcaggtatggatcaccacatctcaatcttggccattcttagtgcagccattgattccaactccttagctttgtttcatgatcacttggcaacaatagcttttttttctttctagataatggaatacaaactacaacttttgcataattttccatgatcgctcggcaccgatagcatttttttctagataaaagaataaaaatcacgacctctacatcatataatgaatggaataaaaatcacggcctttgcatcatataatgcacgcaaccgggcaccaatggcatgcatctcatgtagtcatgtccctatcatttggccatgccctcttcgcatgcacattcatgttcctagtaataactaatctctgtagaacaatagaagagcggaacagtagatgaattatgcgttgattgacagaataattgtgggggggtatatagatcgggtacgtacatgaggatttgaagatcaagccgaatctttccagagataaaaaaaaaagaaaaaaattatgtcagataataactggtcctaatacgtcactagggatagcatcacgaggctttctcccatgcttatacctcctctgttatcatcttccttgagggagtgatcttactgtgcttaaggaggtatctaaagcggtggtgaagccagcgacatcgtcgagcccggataactaatagtgttttgtagtgatcaatccatttttacacgcgtcaacggtgattggtagtggagacgccgacagcgaagcctagacggtcgttcgacctggagccgtctccacctctgatcgaaagatatagtatggtattttcgcctctcacactgggagatacaactttcggcctacaaaatcaacttctggttttaaatctaagatatagattatttctagattcaagcttagaaattgttttttttaagagacagagatagtaccaaattctatttctataattttcttaaatacagtaaattatttcttctcgcgaggaggattacactgcgtaggaacactaggtcatttttcatgtgaaagaaaaaaatggaagctacgaggaagacaaataatttcggcgcgcggggcgaaaaagaagaagatcaaggaagatacccaaggtgggaggtcggtccagcctctattgcgaatagagcggggtgttgttccgcgcgtggggccacacgtgcgcgcacatgcggcgcatatccatcccttcccccccacacccacatgtgtgcatgcatgcggactacgccgctgtacgcgtgcgtggcttacggcgcccgctgctacagctacagcgcccgcacgtgtcagctaccacgccacgccctgcatgcccaccgccgctgcacctccccttgagacgtggtcgctgggcggtggtgggtctcacatgtcagtacgtctttcgtcaaaaaaaaaattcaactttaatttgtttttttcatggtctcacaaccatgtagcttaccaatgcaaacagactaatctaaaaaaaaaaaaataagcttagctataatgcacgcttcgattaagcaatgttacgatgcgtgtgcttagctcaggatagtgcctatattgacacgaaaatatactacatccatataaaaaaatataagcattttttactataaatttatatatttattcatatttatagctaaaaatacttatattttgggtcggagcgagtatcatattttgattaccagtcgtccatcagctatggagtctgtatagggctacgtcctacgttcgggaggtagaaaatatggtgatttattttatttttttgcggggaaatatggtgatttattaatatatgattaattaagtattagttaaaaacttaaagaataggttaatataaatttaaaaataatttttatataaaaaatcgtattgtttagtagtttagaaaatgtgccggatcgggttgagttgggaaaggaggcgggaagaagcgtggtgtagtaattggcgaagggggtaagaagggggagaccggagacatgggtgtgatggttaggcagccggaatgggctctctatcgactcgaacagttggatgagatgagacgagatgggtgacatgcgtgcgtgcatgcatactttgtgccttgtccggtccgtgactgaactgaatcccaggaacgcgaatataatactgtagcacgcacgcacgcacgcacgcatccagctgatctagccggccgagatgtgccactttgaccggtcgtcagacacgctgcatcaatgtaggagcacgtgccagggtatggtcgctgcatgcctgcctgcaaacaatactatactcctcccataaacaaatcaaactccaacaagagaatccgacttgctagaaaaacatcatatcactttaaacaatggagattggagagtccatcagtccaatgtgtttttctgtcttctttgaccatcttccaattccccattatgcttgtgctgacctctatctaattgatcctaagagaagcgttttgcaacatcttgtaaaaaagcaaatcacataatattactcgtgctttatgtagctaagctacattggatcaggatcctctgcagtcaaagtcacgtgactttgtcactgacattgtgggtccgataatcattgggtccacatgtcttacaaagacaatccggctacatcatcatgccagtttccgtaataaattgaccagtaatttagtactatggactttagaacctggcacatgcatagcacagattcatgcatgcgtgcactgcatctgcacgcatttgcattggttgcacaagcagctagctggcctacatataacatattactgtgttatgcgagcaactggccactgtccacatgaagccagtaaatcaactatcaacattgatagtagtaaagctcttgttcagagttgcaggcatataggtgctccctgcatggtttgctcaaacttaagatagatgatactacattctaaaaatgcaagtatttctggtatgtatctggacaagatggttaggactctagctataacatagaaaatgcttatattagaatacacttcattcgtctcaaaatataaggtattttgatcggatatcgatcccatccggttaaataccttaagtagggagtggcagtggcatcaccatcgatcatgtgttattgataaatgattatgattatgagatctcttatctacaatctctttctgaataatcgacctgctgatccatccattgcatgcatggttgttgcctgacaaggacattgaaatggactgctaattgctgccatggatgcaggtgggtggcggaggaggaccacatcgtcaacttccccaccgtgcggctcaagcggggcatgcccatcagggtcaccgccaaggaggacgacgactag</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001048832.1 RefSeq:Os01g0197100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 1]]
[[Category:Chromosome 1]]

Os05g0276500

2014-05-24T08:52:25Z

LQ: /* Annotated Information */

Please input one-sentence summary here.

==Annotated Information==
Os05g0276500 locate at the chromosome5,it's gene type is protein coding,and expression information at root and stem.
===Function===
Please input function information here.

===Expression===
Please input expression information here.

===Evolution===
Please input evolution information here.

You can also add sub-section(s) at will.

==Labs working on this gene==
Please input related labs here.

==References==
Please input cited references here.

==Structured Information==
{{JaponicaGene|
GeneName = Os05g0276500|
Description = Expansin Os-EXPA3|
Version = NM_001061612.1 GI:115462954 GeneID:4338278|
Length = 1781 bp|
Definition = Oryza sativa Japonica Group Os05g0276500, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 5|Chromosome 5]]|
AP = Chromosome 5:11404176..11405956|
CDS = 11404443..11404752,11405334..11405655,11405755..11405890|
GCID = <gbrowseImage1>
name=NC_008398:11404176..11405956
source=RiceChromosome05
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008398:11404176..11405956
source=RiceChromosome05
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgctctctggaatggagaaacaacctgcaatgcttctcgttcttgtgacactctgcgccttcgcatgcaagcgatccgtcgcacagtctgcctttgccacgttctacggtggcaaggacgggtcgggcaccatgggtggcgcgtgtgggtacggtaacctgtacaacgccgggtacgggctgtacaacgcggcgctgagctcggcgctgttcaacgacggcgccatgtgcggcgcgtgctacaccatcacctgcgacaccagccagaccaagtggtgcaagcccggcggcaactccatcaccatcacggccaccaacttatgcccgcctaactgggcgctgcccagcaacagcggcgggtggtgcaacccaccgcggcagcacttcgacatgtcacagccggcctgggagaacatcgccgtctatcaagccggcattgtccccgtcaactacaagagggttccgtgccagaggagcggggggatccggttcgcgatcagcggtcacgactactttgagcttgtgaccgtgacgaacgtgggcggcagcggcgtggtggcgcagatgtcgatcaaggggtccaacacgggctggatggcgatgagcaggaattggggcgccaattggcagagcaacgcgtacctcgccgggcagagcctgtccttcattgtgcagctcgacgacggccgaaaggtcacggcctggaacgtcgccccgtccaactggttcttcggtgccacctactctacctcttgggtgcagttctga</cdnaseq>|
AA = <aaseq>MLSGMEKQPAMLLVLVTLCAFACKRSVAQSAFATFYGGKDGSGT MGGACGYGNLYNAGYGLYNAALSSALFNDGAMCGACYTITCDTSQTKWCKPGGNSITI TATNLCPPNWALPSNSGGWCNPPRQHFDMSQPAWENIAVYQAGIVPVNYKRVPCQRSG GIRFAISGHDYFELVTVTNVGGSGVVAQMSIKGSNTGWMAMSRNWGANWQSNAYLAGQ SLSFIVQLDDGRKVTAWNVAPSNWFFGATYSTSWVQF</aaseq>|
DNA = <dnaseqindica>1205..1514#302..623#67..202#acaaaacgaaaacctaaagaatcaccagccttgcaaaccaatcaacctgtctagattaatttttgcatgctctctggaatggagaaacaacctgcaatgcttctcgttcttgtgacactctgcgccttcgcatgcaagcgatccgtcgcacagtctgcctttgccacgttctacggtggcaaggacgggtcgggcaccatgggtacgtacatttcaagcttaatttgatattttctcaaacaaattagctgatgatgatgatgccatttctgaacttggatgtacgtgcgacgcgatgcaggtggcgcgtgtgggtacggtaacctgtacaacgccgggtacgggctgtacaacgcggcgctgagctcggcgctgttcaacgacggcgccatgtgcggcgcgtgctacaccatcacctgcgacaccagccagaccaagtggtgcaagcccggcggcaactccatcaccatcacggccaccaacttatgcccgcctaactgggcgctgcccagcaacagcggcgggtggtgcaacccaccgcggcagcacttcgacatgtcacagccggcctgggagaacatcgccgtctatcaagccggcattgtccccgtcaactacaagaggtacatgcatcctactcctagttgaataatagccgaaatcgttatagatcaaagatcaatcggagatatatatatgtttgaagtgtttttttttactcgaattgatatacacactcactagctagctcatgaaggcacatcgataacgggtctagaaggattataggttcatgcttgtttcatgctagcaccaatttttcagtaccaatgctatcgatcactagttctagttttgaaacggcttatatgtatcgatactagtgtaaaaccagtacctaccagactaccagagatgggctaacagactcaaagcaggtactgtaatagactataaatcggctacaaacttatgtagagtagataagagaggagagataagagaagcggactacttagccagctgcagcacgtacatagatactgacgatgatttggagctagtaatttactatactattaaacttgctctcataattaccggtgctagactagaaacggcacctataatctgacataggtgcacacagctagtagtaaggtttctcaaaaaacctcacaaatgctgcttgtgtccaatgcagggttccgtgccagaggagcggggggatccggttcgcgatcagcggtcacgactactttgagcttgtgaccgtgacgaacgtgggcggcagcggcgtggtggcgcagatgtcgatcaaggggtccaacacgggctggatggcgatgagcaggaattggggcgccaattggcagagcaacgcgtacctcgccgggcagagcctgtccttcattgtgcagctcgacgacggccgaaaggtcacggcctggaacgtcgccccgtccaactggttcttcggtgccacctactctacctcttgggtgcagttctgaagtactgcaacgtacgtacaggtttagttgataaattatacacatgctgttcatggacataattaattaattactagccatcccggccgtttgtgcatgttctgctaggttgtgtgttgcgtggcattaacttgtttttctctgttaccagtcgttttgtcaaacgcgttggttgtggcaagcgcgattgaatgttttgcccaccaattaatgtaactgatcgtatgggtgttgagttaataaattatgtttaattatgcggatgcc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001061612.1 RefSeq:Os05g0276500]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 5]]
[[Category:Chromosome 5]]

Os05g0276500

2014-05-24T08:50:49Z

LQ: /* Annotated Information */

Please input one-sentence summary here.

==Annotated Information==
===Function===
Please input function information here.

===Expression===
Please input expression information here.

===Evolution===
Please input evolution information here.

You can also add sub-section(s) at will.
Os05g0276500 locate at the chromosome5,it's gene type is protein coding,and expression information at root and stem.

==Labs working on this gene==
Please input related labs here.

==References==
Please input cited references here.

==Structured Information==
{{JaponicaGene|
GeneName = Os05g0276500|
Description = Expansin Os-EXPA3|
Version = NM_001061612.1 GI:115462954 GeneID:4338278|
Length = 1781 bp|
Definition = Oryza sativa Japonica Group Os05g0276500, complete gene.|
Source = Oryza sativa Japonica Group

ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
|
Chromosome = [[:category:Japonica Chromosome 5|Chromosome 5]]|
AP = Chromosome 5:11404176..11405956|
CDS = 11404443..11404752,11405334..11405655,11405755..11405890|
GCID = <gbrowseImage1>
name=NC_008398:11404176..11405956
source=RiceChromosome05
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008398:11404176..11405956
source=RiceChromosome05
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgctctctggaatggagaaacaacctgcaatgcttctcgttcttgtgacactctgcgccttcgcatgcaagcgatccgtcgcacagtctgcctttgccacgttctacggtggcaaggacgggtcgggcaccatgggtggcgcgtgtgggtacggtaacctgtacaacgccgggtacgggctgtacaacgcggcgctgagctcggcgctgttcaacgacggcgccatgtgcggcgcgtgctacaccatcacctgcgacaccagccagaccaagtggtgcaagcccggcggcaactccatcaccatcacggccaccaacttatgcccgcctaactgggcgctgcccagcaacagcggcgggtggtgcaacccaccgcggcagcacttcgacatgtcacagccggcctgggagaacatcgccgtctatcaagccggcattgtccccgtcaactacaagagggttccgtgccagaggagcggggggatccggttcgcgatcagcggtcacgactactttgagcttgtgaccgtgacgaacgtgggcggcagcggcgtggtggcgcagatgtcgatcaaggggtccaacacgggctggatggcgatgagcaggaattggggcgccaattggcagagcaacgcgtacctcgccgggcagagcctgtccttcattgtgcagctcgacgacggccgaaaggtcacggcctggaacgtcgccccgtccaactggttcttcggtgccacctactctacctcttgggtgcagttctga</cdnaseq>|
AA = <aaseq>MLSGMEKQPAMLLVLVTLCAFACKRSVAQSAFATFYGGKDGSGT MGGACGYGNLYNAGYGLYNAALSSALFNDGAMCGACYTITCDTSQTKWCKPGGNSITI TATNLCPPNWALPSNSGGWCNPPRQHFDMSQPAWENIAVYQAGIVPVNYKRVPCQRSG GIRFAISGHDYFELVTVTNVGGSGVVAQMSIKGSNTGWMAMSRNWGANWQSNAYLAGQ SLSFIVQLDDGRKVTAWNVAPSNWFFGATYSTSWVQF</aaseq>|
DNA = <dnaseqindica>1205..1514#302..623#67..202#acaaaacgaaaacctaaagaatcaccagccttgcaaaccaatcaacctgtctagattaatttttgcatgctctctggaatggagaaacaacctgcaatgcttctcgttcttgtgacactctgcgccttcgcatgcaagcgatccgtcgcacagtctgcctttgccacgttctacggtggcaaggacgggtcgggcaccatgggtacgtacatttcaagcttaatttgatattttctcaaacaaattagctgatgatgatgatgccatttctgaacttggatgtacgtgcgacgcgatgcaggtggcgcgtgtgggtacggtaacctgtacaacgccgggtacgggctgtacaacgcggcgctgagctcggcgctgttcaacgacggcgccatgtgcggcgcgtgctacaccatcacctgcgacaccagccagaccaagtggtgcaagcccggcggcaactccatcaccatcacggccaccaacttatgcccgcctaactgggcgctgcccagcaacagcggcgggtggtgcaacccaccgcggcagcacttcgacatgtcacagccggcctgggagaacatcgccgtctatcaagccggcattgtccccgtcaactacaagaggtacatgcatcctactcctagttgaataatagccgaaatcgttatagatcaaagatcaatcggagatatatatatgtttgaagtgtttttttttactcgaattgatatacacactcactagctagctcatgaaggcacatcgataacgggtctagaaggattataggttcatgcttgtttcatgctagcaccaatttttcagtaccaatgctatcgatcactagttctagttttgaaacggcttatatgtatcgatactagtgtaaaaccagtacctaccagactaccagagatgggctaacagactcaaagcaggtactgtaatagactataaatcggctacaaacttatgtagagtagataagagaggagagataagagaagcggactacttagccagctgcagcacgtacatagatactgacgatgatttggagctagtaatttactatactattaaacttgctctcataattaccggtgctagactagaaacggcacctataatctgacataggtgcacacagctagtagtaaggtttctcaaaaaacctcacaaatgctgcttgtgtccaatgcagggttccgtgccagaggagcggggggatccggttcgcgatcagcggtcacgactactttgagcttgtgaccgtgacgaacgtgggcggcagcggcgtggtggcgcagatgtcgatcaaggggtccaacacgggctggatggcgatgagcaggaattggggcgccaattggcagagcaacgcgtacctcgccgggcagagcctgtccttcattgtgcagctcgacgacggccgaaaggtcacggcctggaacgtcgccccgtccaactggttcttcggtgccacctactctacctcttgggtgcagttctgaagtactgcaacgtacgtacaggtttagttgataaattatacacatgctgttcatggacataattaattaattactagccatcccggccgtttgtgcatgttctgctaggttgtgtgttgcgtggcattaacttgtttttctctgttaccagtcgttttgtcaaacgcgttggttgtggcaagcgcgattgaatgttttgcccaccaattaatgtaactgatcgtatgggtgttgagttaataaattatgtttaattatgcggatgcc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001061612.1 RefSeq:Os05g0276500]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 5]]
[[Category:Chromosome 5]]