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| + | The rice '''''Os06g0660200''''' was reported as '''''OsPIN2''''' in 2012 <ref name="ref1" /> by researchers from China. |
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| | ==Annotated Information== | | ==Annotated Information== |
| | ===Function=== | | ===Function=== |
| − | OsPIN2was proposed as an auxin efflux transporter and a member of the PIN family (Wanget al., 2009)<ref name="pmid:19825657" />. The cDNA
| + | * Crop architecture parameters such as tiller number, angle and plant height are important agronomic traits that have been considered for breeding programmes. Auxin distribution within the plant has long been recognized to alter architecture. |
| − | sequence of OsPIN2(AK101191) comprises 2276 nucleotides and alignment between the cDNA, and the genomic sequence
| + | * '''''OsPIN2''''' has a distinct auxin-dependent regulation pathway together with '''''OsPIN1b''''' and '''''OsTAC1''''' controlling rice shoot architecture. |
| − | revealed that the gene has seven exons and six introns (Figure S1A). The ORF encodes a protein of 629 amino acid
| + | * Altering '''''OsPIN2''''' expression by genetic transformation can be directly used for modifying rice architecture. |
| − | residues and shows 56.3% sequence identity with Arabidopsis AtPIN2 .Hydropathy and transmembrane (TM) motif
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| − | analyses of the amino acid sequence revealed that the OsPIN2 protein contains three characteristic regions, including a
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| − | hydrophobic region with five TM domains, a predominantly hydrophilic core, followed by another hydrophobic region with
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| − | four TM segments .
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| − | '''The gain of function ofOsPIN2in rice plants''' | + | ===Phenotypic analysis=== |
| − | | + | * Over-expression of '''''OsPIN2''''' through a transgenic approach in rice (Japonica cv. Nipponbare) led to a shorter plant height, more tillers and a larger tiller angle when compared with wild type (WT). |
| − | It was previously reported that knockdown of OsPIN1bexpression very significantly increased the number of tillers and tiller
| + | * Over-expression of OsPIN2 enhanced auxin transport from shoots to roots, but did not alter the polar auxin pattern in the roots. |
| − | angle, while over-expression did not alter the shoot architecture
| + | * Transgenic plants were less sensitive to N- 1 -naphthylphtha-lamic acid, an auxin transport inhibitor, than WT in their root growth. |
| − | of rice (Xuet al., 2005)<ref name="pmid:16085936" />. In the rice genome, OsLazy1 has been
| + | * OsPIN2-over-expressing plants had suppressed the expression of a gravitropism-related gene OsLazy1 in the shoots, but unaltered expression of OsPIN1b and OsTAC1, which were reported as tiller angle controllers in rice. |
| − | identified as a novel grass-specific protein playing a negative
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| − | role in polar auxin transport; its mutation resulted in an increase
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| − | in the rice tiller angle (Abe et al., 1996<ref name="pmid:11539861" />; Li et al., 2007<ref name="pmid:17468779" />; Yoshihara and Iino, 2007<ref name="pmid:17412736" /> ). It is interesting to speculate whether
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| − | enhancingOsLazy1expression could compact rice architecture
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| − | further. In addition, OsTAC1was defined as a tiller angle
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| − | control gene in rice, its expression level positively correlated
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| − | with tiller angle and numbers (Yuet al., 2007))<ref name="pmid:17908158" />. | |
| − | Transgenic plants over-expressingOsPIN2showed larger tiller | |
| − | angles, lower plant height, more tillers than WT (Figure 2c–e),
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| − | which was similar to theOsPIN1bknockdown mutants (Xu
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| − | et al., 2005)<ref name="pmid:16085936" />. The change of phenotype caused by alteration of
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| − | OsPIN2andOsPIN1bexpression supported the common observation that rice plant height was negatively correlated with tiller
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| − | number (Wang and Li, 2005). Therefore, auxin might play a key
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| − | role in the crosstalk between the plant height and branching,
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| − | an aspect of canopy architecture which is poorly understood.
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| | | | |
| | ===Expression=== | | ===Expression=== |
| − | '''Generation of transgenic rice showingOsPIN2 over-expression and phenotypes of transformants'''
| + | * The expression patterns of the auxin reporter DR5::GUS and quantification of auxin distribution showed that '''''OsPIN2''''' over-expression increased auxin transport from the shoot to the root–shoot junction, resulting in a non-tissue-specific accumulation of more free auxin at the root–shoot junction relative to WT. |
| − | Ubiquitin promoter (pUbi) has been long reported as a useful
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| − | strong promoter in a variety of applications in gene transfer
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| − | studies and drives the gene expression most actively in rapidly
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| − | dividing cells (Cornejoet al., 1993)<ref name="pmid:8219091" /> . To investigate the function
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| − | ofOsPIN2in rice, we made transgenic rice plants over-expressing the full length of cDNA driven by pUbi. Southern blotting
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| − | analysis revealed that they came from two transgenic lines (O1
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| − | and O2, respectively) distinct from each other and both had | |
| − | one copy of the transgene (Figure 1a). Real-time RT-PCR
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| − | analyses showed that transcriptional expression ofOsPIN2was
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| − | moderate in the root–shoot junctions and roots but very faint in
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| − | the leaves of WT (the untransformed plants as control)
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| − | (Figure 1b). In both the transgenic lines, OsPIN2had the most
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| − | abundant transcripts in the root–shoot junction, and similar
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| − | levels in the leaves and roots (Figure 1b). The increases in
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| − | OsPIN2transcriptional expression driven by pUbi in these tissues
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| − | reached by 3–15 times.
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| − | | |
| − | Over-expression ofOsPIN2significantly increased tiller angle
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| − | and numbers and decreased height of the rice in comparison
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| − | with WT (Figure 1i). At 80 days after germination,
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| − | O1 and O2 plants had a wider tiller angle (the angle between
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| − | the outermost tillers in the left and right side) (44° and 48°,
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| − | respectively) than the WT (22°). In addition, the two
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| − | transgenic lines had many more tillers per plant (21 and 23,
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| − | respectively) than the WT (17 on the average). In
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| − | contrast, the whole plant height was very significantly
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| − | decreased byOsPIN2over-expression. The transgenic
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| − | plant phenotypes were inheritable from T0 (Figure 2f–h) to T2
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| − | generations.
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| − | At the ripening stage, theOsPIN2-over-expressing
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| − | plants showed shorter panicle length, less number of grains per
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| − | panicle and lower grain weight per panicle in comparison with
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| − | the control. Even though the seed setting rate was
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| − | similar between O1, O2 and WT, O1 plants had a 35% increase
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| − | in the effective tiller number, leading to a 16% grain yield
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| − | increase per plant relative to control plants. O2 plants
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| − | showed the same tendency but not significant increase in panicle number and grain yield. OsPIN2over-expression
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| − | also decreased the grain length and breadth by 4%–7% and
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| − | 12%–16%, respectively. The changes in the grain size
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| − | lead to 10%–12.5% decreases of 1000-grain weight compared
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| − | to WT. Moreover, over-expression of the OsPIN2significantly
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| − | decreased the number of adventitious roots and the total root
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| − | length by 22%–28%.
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| − | | |
| − | | |
| − | | |
| − | There was no putative gene in the insertion site for both O1 and O2. This direct genetic evidence further supported the OsPIN2 contribution to the altered
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| − | phenotype of the transgenic rice in Figure 1
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| − | | |
| − | [[File:Phenotypes and molecular analyses ofOsPIN2over-expression transgenic plants (O1 and O2) and untranformed wild-type plant (WT).jpg]]
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| − | | |
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| − | | |
| − | ===Evolution===
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| − | '''Structure and evolution of the PIN super family''' | |
| − | | |
| − | Genes homologous to the Arabidopsis PIN sequences are present in genomes throughout the plant kingdom. These include representatives in non-vascular plants such as Physcomitrella; in vascular plants, PIN sequences appear to be ubiquitous. Both before and since the divergence of monocot and dicot plants there have been significant changes in the number and the structure of PINs. However, in dicot plants, the phylogenetic structure of the family has been broadly conserved ( Figure 2a). Both Medicago and potato contain at least five PIN sequences (these data were collected from EST sequencing projects and, accordingly, have the propensity to exclude sequences from mRNA expressed either at a low level or exclusively in certain tissue types). Each of these genes shows similarity to one of the eight PINs of Arabidopsis. Both Medicago and potato contain two genes homologous to AtPIN1 and two genes homologous to the AtPIN3, AtPIN4, AtPIN7 group. Medicago has one gene homologous to AtPIN2. Potato has one gene homologous to AtPIN6. There is additional evidence for such a broadly conserved phylogenetic structure in the PIN genes of soybean (Glycine max). Here, there exist single homologous genes to AtPIN1, AtPIN2, AtPIN5, AtPIN8, and two genes homologous to the AtPIN3, AtPIN4, AtPIN7 cluster of Arabidopsis.
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| − | | |
| − | [[File:Neighbour-joining phylogenetic trees showing the predicted relationship between Arabidopsis AtPINs and (a) dicot (Medicago, soybean and potato) and (b) monocot (rice and wheat) PINs.jpg]]
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| − | | |
| − | To confirm that the changed phenotype of O1 and O2 was
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| − | caused by over-expression of OsPIN2in the transgenic plants,
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| − | we detected the location of one copy T-DNA insertion in these
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| − | two lines. The T-DNA containing ubi-promoter was inserted in
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| − | the chromosome 3 and 10 in the genome of O1 and O2,
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| − | respectively (Figure 2).
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| − | | |
| − | [[File:Schematic map of integrated T-DNA insertion site in rice genome.jpg]]
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| − | | |
| − | ==Labs working on this gene==
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| − | 1.State Key Laboratory of Crop Genetics and Germplasm Enhancement, College of Resources and Environmental Sciences, Nanjing Agricultural University, Nanjing,
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| − | China
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| − | | |
| − | 2.Institut für Biologie II, Universität Freiburg, Schänzlestrasse 1, 79104 Freiburg, Germany.
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| | ==References== | | ==References== |
| | <references> | | <references> |
| − | <ref name="pmid:19825657"> Wang, J.R., Hu, H., Wang, G.H., Li, J., Chen, J.Y. and Wu, P. (2009) | + | * <ref name="ref1"> Chen Y, Fan X, Song W, Zhang Y, Xu G. Over-expression of OsPIN2 leads to |
| − | Expression of PIN Genes in Rice (Oryza sativaL.): tissue specificity and
| + | increased tiller numbers, angle and shorter plant height through suppression of |
| − | regulation by hormones. Mol. Plant, 2, 823–831. </ref>
| + | OsLAZY1. Plant Biotechnol J. 2012 Feb;10(2):139-49. doi: |
| − | <ref name="pmid:16085936"> Xu, M., Zhu, L., Shou, H.X. and Wu, P. (2005) A PIN1 family gene, OsPIN1,
| + | 10.1111/j.1467-7652.2011.00637.x. PubMed PMID: 21777365. |
| − | involved in auxin-dependent adventitious root emergence and tillering in
| + | </ref> |
| − | rice. Plant Cell Physiol.46, 1674–1681. </ref>
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| − | <ref name="pmid:17908158"> Yu, B., Lin, Z., Li, H., Li, X., Li, J., Wang, Y., Zhang, X., Zhu, Z., Zhai, W.,
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| − | Wang, X., Xie, D. and Sun, C. (2007) TAC1, a major quantitative trait
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| − | locus controlling tiller angle in rice. Plant J.52, 891–898.</ref>
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| − | <ref name="pmid:11539861"> Abe, K., Takahashi, H. and Suge, H. (1996) Lazy gene (la) responsible for
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| − | both an agravitropism of seedlings and lazy habit of tiller growth in rice
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| − | (Oryza sativaL).J. Plant. Res.109, 381–386. </ref>
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| − | <ref name="pmid:17468779"> Li, P., Wang, Y., Qian, Q., Fu, Z., Wang, M., Zeng, D., Li, B., Wang, X. and
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| − | Li, J. (2007) LAZY1 controls rice shoot gravitropism through regulating
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| − | polar auxin transport.Cell Res.17, 402–410.</ref>
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| − | <ref name="pmid:17412736"> Yoshihara, T. and Iino, M. (2007) Identification of the gravitropism-related
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| − | rice gene LAZY1 and elucidation of LAZY1-dependent and -independent
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| − | gravity signaling pathways.Plant Cell Physiol.48, 678–688 </ref>
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| − | <ref name="pmid:8219091"> Cornejo, M.J., Luth, D., Blankenship, K.M., Anderson, O.D. and Blechl, A.E.
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| − | (1993) Activity of a maize ubiquitin promoter in transgenic rice. Plant Mol.
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| − | Biol.23, 567–581.</ref>
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| − | | |
| − | | |
| − | | |
| | </references> | | </references> |
| | | | |
| | ==Structured Information== | | ==Structured Information== |
| − | {{JaponicaGene|
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| − | GeneName = Os06g0660200|
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| − | Description = Similar to Auxin efflux carrier protein|
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| − | Version = NM_001064803.1 GI:115469337 GeneID:4341736|
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| − | Length = 3804 bp|
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| − | Definition = Oryza sativa Japonica Group Os06g0660200, complete gene.|
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| − | Source = Oryza sativa Japonica Group
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| | | | |
| − | ORGANISM Oryza sativa Japonica Group
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| − | Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
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| − | Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
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| − | clade; Ehrhartoideae; Oryzeae; Oryza.
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| − | |
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| − | Chromosome = [[:category:Japonica Chromosome 6|Chromosome 6]]|
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| − | AP = Chromosome 6:28077811..28081614|
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| − | CDS = 28077886..28078901,28078996..28079180,28079269..28079572,28080611..28080696,28080796..28080953<br>,28081075..28081151,28081241..28081307|
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| − | GCID = <gbrowseImage1>
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| − | name=NC_008399:28077811..28081614
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| − | source=RiceChromosome06
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| − | preset=GeneLocation
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| − | </gbrowseImage1>|
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| − | GSID = <gbrowseImage2>
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| − | name=NC_008399:28077811..28081614
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| − | source=RiceChromosome06
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| − | preset=GeneLocation
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| − | </gbrowseImage2>|
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| − | CDNA = <cdnaseq>atgatcaccggacgcgacatctacgacgtgctggcggcgatcgtgccgctgtacgtggcgatgttcctggcgtacgggtcggtgcggtggtgggggatattcacgccggaccagtgctccggcatcaaccgcttcgtcgccgtcttcgccgtgccgctcctctccttccacttcatctccaccaacgacccctactccatgaactaccgcttcctcgccgccgactcgctccagaagctcgtcatcctcgccgcgctcgccgtctggcacaacctgctctcccgctaccgccgcaatggcggcgccgccgcgtcgctcgactggaccatcaccctcttctcgctgtccacgctgcccaacacgctggtcatgggcatcccgctgctccgcgccatgtacggcgacttctccggctcgctcatggtgcagatcgtcgtgctccagagcgtcatctggtacaccctcatgctcttcctcttcgagtaccgcggcgccaaggcgctcatctccgagcagttcccgccggacgtcggcgccagcatcgcctcgttccgcgtcgactccgacgtcgtctcgctcaacgggagggaggcgctgcaggcggacgccgaggtggggcgcgacggccgcgtccacgtcgtcatccgccgctccgcctcggcctccaccacgggcggcggcggcggcgcggcgcgctccggcgtgtcccgggcgtacggcgcgtccaacgccatgacgccgcgcgcctccaacctcaccggcgtggagatctactcgctgcagacgtcgcgcgagcccacgccgcgggcgtccagcttcaaccaggccgacttctacgccatgttctccggcagcaagatggccagccagatggctagccccatggcgcagcacggcggcgccggcggccgcgcccagggcctcgacgagcaggtcaccaacaagttcgcctccggcaaggccgccgacccgccgtcgtatcccgccccgaaccccggcatgatgccggcgccaaggaagaaggagctcgggggctcaaactccaactccaacaaggagctacacatgttcgtgtggagctctagcgcgtcgccggtgtcggaggccaacctccgcaacgccgtcaaccacgccgcctccaccgacttcgcctccgcgccgccgccggcagccgttcccgtcggcggcgccactcccaaaggggtgagtggcagtgtcacgccggcggcgaagaacggcggcggcgagttggagatcgaggacgggctgaagagcccggcggcggggctggcggcgaagttcccggtgtcggggtcgccgtacgtggcgccgaggaagaagggcggcggcgccgacgtgcccgggctggcggaggcggcgcacccgatgccgccgacgagcgtgatgacgcggctcatcctcatcatggtgtggcgcaagctcatcagaaaccccaacacctactccagcctcatcggcctcgtctggtccctcgtctccttcaggtggaatatccaaatgccttcaataataaagggctcaatatcaatattgtcagatgcagggctaggaatggctatgttcagcttaggcttgttcatggctctgcaaccaaagatcatttcttgtggcaagaccgttgcgacatttgcaatggcagtgaggttcttgactggtccagctgttattgcagctacttccattgccattgggctcaggggagtactcttgcatgttgccattgttcaggcagcacttccacaaggcattgtcccgtttgtgtttgccaaggagtacaattgccatcctcaaatacttagcacagcggttatttttgggatgctcatcgcgcttccgatcacgatactctactatgtgcttcttgggatatag</cdnaseq>|
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| − | AA = <aaseq>MITGRDIYDVLAAIVPLYVAMFLAYGSVRWWGIFTPDQCSGINR FVAVFAVPLLSFHFISTNDPYSMNYRFLAADSLQKLVILAALAVWHNLLSRYRRNGGA AASLDWTITLFSLSTLPNTLVMGIPLLRAMYGDFSGSLMVQIVVLQSVIWYTLMLFLF EYRGAKALISEQFPPDVGASIASFRVDSDVVSLNGREALQADAEVGRDGRVHVVIRRS ASASTTGGGGGAARSGVSRAYGASNAMTPRASNLTGVEIYSLQTSREPTPRASSFNQA DFYAMFSGSKMASQMASPMAQHGGAGGRAQGLDEQVTNKFASGKAADPPSYPAPNPGM MPAPRKKELGGSNSNSNKELHMFVWSSSASPVSEANLRNAVNHAASTDFASAPPPAAV PVGGATPKGVSGSVTPAAKNGGGELEIEDGLKSPAAGLAAKFPVSGSPYVAPRKKGGG ADVPGLAEAAHPMPPTSVMTRLILIMVWRKLIRNPNTYSSLIGLVWSLVSFRWNIQMP SIIKGSISILSDAGLGMAMFSLGLFMALQPKIISCGKTVATFAMAVRFLTGPAVIAAT SIAIGLRGVLLHVAIVQAALPQGIVPFVFAKEYNCHPQILSTAVIFGMLIALPITILY YVLLGI</aaseq>|
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| − | DNA = <dnaseqindica>76..1091#1186..1370#1459..1762#2801..2886#2986..3143#3265..3341#3431..3497#aattagctcgttctcttgtgtcaagaaaaaaaaagaaagaaaaagctcgccgccgccgccaccgtcgccggcgcgatgatcaccggacgcgacatctacgacgtgctggcggcgatcgtgccgctgtacgtggcgatgttcctggcgtacgggtcggtgcggtggtgggggatattcacgccggaccagtgctccggcatcaaccgcttcgtcgccgtcttcgccgtgccgctcctctccttccacttcatctccaccaacgacccctactccatgaactaccgcttcctcgccgccgactcgctccagaagctcgtcatcctcgccgcgctcgccgtctggcacaacctgctctcccgctaccgccgcaatggcggcgccgccgcgtcgctcgactggaccatcaccctcttctcgctgtccacgctgcccaacacgctggtcatgggcatcccgctgctccgcgccatgtacggcgacttctccggctcgctcatggtgcagatcgtcgtgctccagagcgtcatctggtacaccctcatgctcttcctcttcgagtaccgcggcgccaaggcgctcatctccgagcagttcccgccggacgtcggcgccagcatcgcctcgttccgcgtcgactccgacgtcgtctcgctcaacgggagggaggcgctgcaggcggacgccgaggtggggcgcgacggccgcgtccacgtcgtcatccgccgctccgcctcggcctccaccacgggcggcggcggcggcgcggcgcgctccggcgtgtcccgggcgtacggcgcgtccaacgccatgacgccgcgcgcctccaacctcaccggcgtggagatctactcgctgcagacgtcgcgcgagcccacgccgcgggcgtccagcttcaaccaggccgacttctacgccatgttctccggcagcaagatggccagccagatggctagccccatggcgcagcacggcggcgccggcggccgcgcccagggcctcgacgagcaggtcaccaacaagttcgcctccggcaaggccgccgacccgccgtcgtatcccgccccgaaccccggcatgatgccggcgccaaggtaaaatgaaactgattattaacacctcaaaatttctgttcatcgatcgtgttttgatcgagttggatttttgattttgtcgccattgctacaggaagaaggagctcgggggctcaaactccaactccaacaaggagctacacatgttcgtgtggagctctagcgcgtcgccggtgtcggaggccaacctccgcaacgccgtcaaccacgccgcctccaccgacttcgcctccgcgccgccgccggcagccgttcccgtcggcggcgccactcccaaaggtactctctcatccatccattgacgcacacgacgacgatcacaccacatgtgttcaaggcttcgtctaatggtgcgtgcatgccacaggggtgagtggcagtgtcacgccggcggcgaagaacggcggcggcgagttggagatcgaggacgggctgaagagcccggcggcggggctggcggcgaagttcccggtgtcggggtcgccgtacgtggcgccgaggaagaagggcggcggcgccgacgtgcccgggctggcggaggcggcgcacccgatgccgccgacgagcgtgatgacgcggctcatcctcatcatggtgtggcgcaagctcatcagaaaccccaacacctactccagcctcatcggcctcgtctggtccctcgtctccttcaggtattggattctcaccgatgttttcggtactcaagctgacgttctcgcttccacagttccacttcatcgacgcccgctttcccggtcgcttccctctaagatggaacactcttctgccaatacattttgacatcccacagctatgacagatcgcttatcccggttcatcttcagggcaagggcgctttttttcaagttataccgaattatcaaaacagattatttactctccatttcaaaaaagtattttataaaaaaaaatattcttctactacattacttagattacttgttttaagctagttgttaagtttttttttaaattgtttaatttatttgtataatgaaatcgataatctgcatagtaatctaaaccaaataagtaatatacacttgagttcacgatctgatcaatttatgttgttcgaagctacaaaaaggatatatttttgctatgctattcgttgctagattgcttgaaattttaatctaggaagaggcaagatcgatgagaacttacttaacttcacaactggattagtgttttctgaacgcatgtgagagcacatgataattacgttggtctcggatctggtccagcacatgcaaagaacatacacacgtatgcagatgcaggtgcaggtgcaggtcttatgtgagccctggagtttagcagcagagaccaaatctaccgaatcagctgcgattgccgtgtttgtccccatccaagacagacacggcgaattcggctcaactagctagatgatccatcatgtgtacaagaagaagagcaaaaaatgtacaatgatatggatgcatcattgattcattgtactactatcttaccattagatggcacatctagagggagggttgcaagaacagggcatttttagtggggaggagcacatgcctgcaagcatctaaaattgcatacatgcatgatacatctagcattgattctcttgtcatacaatttttttttacattgtgaaaatatatgcttaattgcttattgactttgtgaatttggatcaggtggaatatccaaatgccttcaataataaagggctcaatatcaatattgtcagatgcagggctaggaatggctatgttcagcttaggtacagagttgacctttttgatacatatctatagccataactagtgagttttttctttgctttgaccaagaaattctgacaatagtatatgtaacccaggcttgttcatggctctgcaaccaaagatcatttcttgtggcaagaccgttgcgacatttgcaatggcagtgaggttcttgactggtccagctgttattgcagctacttccattgccattgggctcaggggagtactcttgcatgttgccattgttcaggtaagcagtagattctttatttcttgcaatcatcttaaaagaaaaaaaatattaaattttgtaattatggtgaattggaatgtattgtggtgatcttatagattaattgattgtcaaacaggcagcacttccacaaggcattgtcccgtttgtgtttgccaaggagtacaattgccatcctcaaatacttagcacagcgtaagaaatgcatgcttaaactttctatttgttcgtccatacggttgctgaagtatatagcctaaaatatataaatgtgaaattttcagggttatttttgggatgctcatcgcgcttccgatcacgatactctactatgtgcttcttgggatatagtgttcttgaagaaggcaaaaaagaaagagtagggaaaaaaataggattctaggtttctagaggaaaatgcaaaagaaatatgatatgggctttcttgaagacctgaagaactaccagagctgaagaatagggaaatgagatcaagtaggatcctagctagagagaaatgcaaaggaaagacaccccttgattacaattttttaattttttctgcaactgttttggcatcaaagtaaaggttagggccttgagtatgaagagttcagccgttaatttgacaagttgggttggcggtactaaagattcc</dnaseqindica>|
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| − | Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001064803.1 RefSeq:Os06g0660200]|
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| − | }}
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| | [[Category:Genes]] | | [[Category:Genes]] |
| | [[Category:Japonica mRNA]] | | [[Category:Japonica mRNA]] |
