RiceWiki - User contributions [en]

Os02g0232100

2014-06-04T10:27:02Z

Simon:

HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.
===Mutation===
[[File:Figure111.jpg|right|thumb|300px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref3" />).'']]
60Co γ-ray irradiation was used to induce a sterile mutant from the japonica rice variety Wuxiangjing 9.The mutant showed normal vegetative growth but exhibited complete sterility (Figure 1A and 1B). Cytological observation of anthers showed that almost all pollens were shrunken and inviable (Figure 1C and 1D). When pollinated with wild-type (WT) pollens, the mutant spikelets were unable to set any seeds, suggesting that the female gametes were also sterile. The progenies of the heterozygous plants segregated from normal to sterile phenotype in a 3:1 ratio (fertile, 30; sterile, 10), indicating that a single recessive gene is responsible for the sterile phenotype<ref name="ref3" />.
===Expression（Mutant VS Wild type）===
[[File:Figure222.jpg|right|thumb|300px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure333.jpg|right|thumb|300px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure444.jpg|right|thumb|300px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.
===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.
===Knowledge Extension===

In hei10 mutants, only about 31% chiasmata were maintained. This number is similar to that of mer3 mutants, in which about 28% chiasmata were remained. Previous studies revealed that there are at least two classes of CO which occur in budding yeast and Arabidopsis <ref name="ref7"/><ref name="ref8"/><ref name="ref9"/>. Investigations of mer3 mutants in rice showed that apart from a reduced number of chiasmata, the remaining chiasmata distributed randomly among cells, suggesting that rice might also have at least two kinds of COs: one that appears to be sensitive to interference, whereas the other one is not <ref name="ref10"/>. The close number of residual chiasmata indicates similar contributions of HEI10 and MER3 during CO formation. Considering the close number and similar random distribution of residual chiasmata noted in mer3 and hei10 mutants, it is likely that HEI10 and MER3 may act in the same CO pathway, namely an interference-sensitive pathway.Besides a reduced chiasma frequency, hei10 also showed a precocious segregation of sister chromatids of some univalents at anaphase I.

===Labs working on this gene===
*State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
*Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

===References===
<references>
<ref name="ref1"> Toby GG, Gherraby W, Coleman TR, Golemis EA (2003) A novel RING finger protein, human enhancer of invasion 10, alters mitotic progression through regulation of cyclin B levels. Mol Cell Biol 23: 2109–2122. </ref>
<ref name="ref2"> Singh MK, Nicolas E, Gherraby W, Dadke D, Lessin S, et al. (2007) HEI10 negatively regulates cell invasion by inhibiting cyclin B/Cdk1 and other promotility proteins. Oncogene 26: 4825–4832. </ref>
<ref name="ref3">Wang, K., Wang, M., Tang, D., Shen, Y., Miao, C., Hu, Q., ... & Cheng, Z. (2012).The role of rice HEI10 in the formation of meiotic crossovers. PLoS genetics, 8(7), e1002809. </ref>
<ref name="ref4">de Boer E, Stam P, Dietrich AJ, Pastink A, Heyting C (2006) Two levels of interference in mouse meiotic recombination. Proc Natl Acad Sci U S A 103: 9607–9612. </ref>
<ref name="ref5">Lhuissier FG, Offenberg HH, Wittich PE, Vischer NO, Heyting C (2007) The mismatch repair protein MLH1 marks a subset of strongly interfering crossovers in tomato. Plant Cell 19: 862–876. </ref>
<ref name="ref6">Bhalla N, Wynne DJ, Jantsch V, Dernburg AF (2008) ZHP-3 acts at crossovers to couple meiotic recombination with synaptonemal complex disassembly and bivalent formation in C. elegans. PLoS Genet 4: e1000235. </ref>
<ref name="ref7">Borner GV, Kleckner N, Hunter N (2004) Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell 117: 29–45. </ref>
<ref name="ref8"> Mercier R, Jolivet S, Vezon D, Huppe E, Chelysheva L, et al. (2005) Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not. Curr Biol 15: 692–701. </ref>
<ref name="ref9">Higgins JD, Armstrong SJ, Franklin FC, Jones GH (2004) The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis. Genes & Dev 18: 2557–2570. </ref>
<ref name="ref10">Wang K, Tang D, Wang M, Lu J, Yu H, et al. (2009) MER3 is required for normal meiotic crossover formation, but not for presynaptic alignment in rice. J Cell Sci 122: 2055–2063. </ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

Os02g0232100

2014-06-04T10:25:05Z

Simon:

HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.
===Mutation===
[[File:Figure111.jpg|right|thumb|150px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref3" />).'']]
60Co γ-ray irradiation was used to induce a sterile mutant from the japonica rice variety Wuxiangjing 9.The mutant showed normal vegetative growth but exhibited complete sterility (Figure 1A and 1B). Cytological observation of anthers showed that almost all pollens were shrunken and inviable (Figure 1C and 1D). When pollinated with wild-type (WT) pollens, the mutant spikelets were unable to set any seeds, suggesting that the female gametes were also sterile. The progenies of the heterozygous plants segregated from normal to sterile phenotype in a 3:1 ratio (fertile, 30; sterile, 10), indicating that a single recessive gene is responsible for the sterile phenotype<ref name="ref3" />.
===Expression（Mutant VS Wild type）===
[[File:Figure222.jpg|right|thumb|150px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure333.jpg|right|thumb|150px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure444.jpg|right|thumb|150px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.
===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.
===Knowledge Extension===

In hei10 mutants, only about 31% chiasmata were maintained. This number is similar to that of mer3 mutants, in which about 28% chiasmata were remained. Previous studies revealed that there are at least two classes of CO which occur in budding yeast and Arabidopsis <ref name="ref7"/><ref name="ref8"/><ref name="ref9"/>. Investigations of mer3 mutants in rice showed that apart from a reduced number of chiasmata, the remaining chiasmata distributed randomly among cells, suggesting that rice might also have at least two kinds of COs: one that appears to be sensitive to interference, whereas the other one is not <ref name="ref10"/>. The close number of residual chiasmata indicates similar contributions of HEI10 and MER3 during CO formation. Considering the close number and similar random distribution of residual chiasmata noted in mer3 and hei10 mutants, it is likely that HEI10 and MER3 may act in the same CO pathway, namely an interference-sensitive pathway.Besides a reduced chiasma frequency, hei10 also showed a precocious segregation of sister chromatids of some univalents at anaphase I.

===Labs working on this gene===
*State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
*Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

===References===
<references>
<ref name="ref1"> Toby GG, Gherraby W, Coleman TR, Golemis EA (2003) A novel RING finger protein, human enhancer of invasion 10, alters mitotic progression through regulation of cyclin B levels. Mol Cell Biol 23: 2109–2122. </ref>
<ref name="ref2"> Singh MK, Nicolas E, Gherraby W, Dadke D, Lessin S, et al. (2007) HEI10 negatively regulates cell invasion by inhibiting cyclin B/Cdk1 and other promotility proteins. Oncogene 26: 4825–4832. </ref>
<ref name="ref3">Wang, K., Wang, M., Tang, D., Shen, Y., Miao, C., Hu, Q., ... & Cheng, Z. (2012).The role of rice HEI10 in the formation of meiotic crossovers. PLoS genetics, 8(7), e1002809. </ref>
<ref name="ref4">de Boer E, Stam P, Dietrich AJ, Pastink A, Heyting C (2006) Two levels of interference in mouse meiotic recombination. Proc Natl Acad Sci U S A 103: 9607–9612. </ref>
<ref name="ref5">Lhuissier FG, Offenberg HH, Wittich PE, Vischer NO, Heyting C (2007) The mismatch repair protein MLH1 marks a subset of strongly interfering crossovers in tomato. Plant Cell 19: 862–876. </ref>
<ref name="ref6">Bhalla N, Wynne DJ, Jantsch V, Dernburg AF (2008) ZHP-3 acts at crossovers to couple meiotic recombination with synaptonemal complex disassembly and bivalent formation in C. elegans. PLoS Genet 4: e1000235. </ref>
<ref name="ref7">Borner GV, Kleckner N, Hunter N (2004) Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell 117: 29–45. </ref>
<ref name="ref8"> Mercier R, Jolivet S, Vezon D, Huppe E, Chelysheva L, et al. (2005) Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not. Curr Biol 15: 692–701. </ref>
<ref name="ref9">Higgins JD, Armstrong SJ, Franklin FC, Jones GH (2004) The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis. Genes & Dev 18: 2557–2570. </ref>
<ref name="ref10">Wang K, Tang D, Wang M, Lu J, Yu H, et al. (2009) MER3 is required for normal meiotic crossover formation, but not for presynaptic alignment in rice. J Cell Sci 122: 2055–2063. </ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

Os02g0232100

2014-06-04T10:22:57Z

Simon: /* Expression（Mutant VS Wild type） */

HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.
===Mutation===
[[File:Figure111.jpg|right|thumb|150px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref3" />).'']]
60Co γ-ray irradiation was used to induce a sterile mutant from the japonica rice variety Wuxiangjing 9.The mutant showed normal vegetative growth but exhibited complete sterility (Figure 1A and 1B). Cytological observation of anthers showed that almost all pollens were shrunken and inviable (Figure 1C and 1D). When pollinated with wild-type (WT) pollens, the mutant spikelets were unable to set any seeds, suggesting that the female gametes were also sterile. The progenies of the heterozygous plants segregated from normal to sterile phenotype in a 3:1 ratio (fertile, 30; sterile, 10), indicating that a single recessive gene is responsible for the sterile phenotype<ref name="ref3" />.

===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.
===Knowledge Extension===

In hei10 mutants, only about 31% chiasmata were maintained. This number is similar to that of mer3 mutants, in which about 28% chiasmata were remained. Previous studies revealed that there are at least two classes of CO which occur in budding yeast and Arabidopsis <ref name="ref7"><ref name="ref8"><ref name="ref9">. Investigations of mer3 mutants in rice showed that apart from a reduced number of chiasmata, the remaining chiasmata distributed randomly among cells, suggesting that rice might also have at least two kinds of COs: one that appears to be sensitive to interference, whereas the other one is not <ref name="ref10">. The close number of residual chiasmata indicates similar contributions of HEI10 and MER3 during CO formation. Considering the close number and similar random distribution of residual chiasmata noted in mer3 and hei10 mutants, it is likely that HEI10 and MER3 may act in the same CO pathway, namely an interference-sensitive pathway.Besides a reduced chiasma frequency, hei10 also showed a precocious segregation of sister chromatids of some univalents at anaphase I.

===Labs working on this gene===
*State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
*Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

===References===
<references>
<ref name="ref1"> Toby GG, Gherraby W, Coleman TR, Golemis EA (2003) A novel RING finger protein, human enhancer of invasion 10, alters mitotic progression through regulation of cyclin B levels. Mol Cell Biol 23: 2109–2122. </ref>
<ref name="ref2"> Singh MK, Nicolas E, Gherraby W, Dadke D, Lessin S, et al. (2007) HEI10 negatively regulates cell invasion by inhibiting cyclin B/Cdk1 and other promotility proteins. Oncogene 26: 4825–4832. </ref>
<ref name="ref3">Wang, K., Wang, M., Tang, D., Shen, Y., Miao, C., Hu, Q., ... & Cheng, Z. (2012).The role of rice HEI10 in the formation of meiotic crossovers. PLoS genetics, 8(7), e1002809. </ref>
<ref name="ref4">de Boer E, Stam P, Dietrich AJ, Pastink A, Heyting C (2006) Two levels of interference in mouse meiotic recombination. Proc Natl Acad Sci U S A 103: 9607–9612. </ref>
<ref name="ref5">Lhuissier FG, Offenberg HH, Wittich PE, Vischer NO, Heyting C (2007) The mismatch repair protein MLH1 marks a subset of strongly interfering crossovers in tomato. Plant Cell 19: 862–876. </ref>
<ref name="ref6">Bhalla N, Wynne DJ, Jantsch V, Dernburg AF (2008) ZHP-3 acts at crossovers to couple meiotic recombination with synaptonemal complex disassembly and bivalent formation in C. elegans. PLoS Genet 4: e1000235. </ref>
<ref name="ref7">Borner GV, Kleckner N, Hunter N (2004) Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell 117: 29–45. </ref>
<ref name="ref8"> Mercier R, Jolivet S, Vezon D, Huppe E, Chelysheva L, et al. (2005) Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not. Curr Biol 15: 692–701. </ref>
<ref name="ref9">Higgins JD, Armstrong SJ, Franklin FC, Jones GH (2004) The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis. Genes & Dev 18: 2557–2570. </ref>
<ref name="ref10">Wang K, Tang D, Wang M, Lu J, Yu H, et al. (2009) MER3 is required for normal meiotic crossover formation, but not for presynaptic alignment in rice. J Cell Sci 122: 2055–2063. </ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

Os02g0232100

2014-06-04T10:20:55Z

Simon:

HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.
===Mutation===
[[File:Figure111.jpg|right|thumb|150px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref3" />).'']]
60Co γ-ray irradiation was used to induce a sterile mutant from the japonica rice variety Wuxiangjing 9.The mutant showed normal vegetative growth but exhibited complete sterility (Figure 1A and 1B). Cytological observation of anthers showed that almost all pollens were shrunken and inviable (Figure 1C and 1D). When pollinated with wild-type (WT) pollens, the mutant spikelets were unable to set any seeds, suggesting that the female gametes were also sterile. The progenies of the heterozygous plants segregated from normal to sterile phenotype in a 3:1 ratio (fertile, 30; sterile, 10), indicating that a single recessive gene is responsible for the sterile phenotype<ref name="ref3" />.

===Expression（Mutant VS Wild type）===
[[File:Figure222.jpg|right|thumb|150px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure333.jpg|right|thumb|150px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure444.jpg|right|thumb|150px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.

===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.
===Knowledge Extension===

In hei10 mutants, only about 31% chiasmata were maintained. This number is similar to that of mer3 mutants, in which about 28% chiasmata were remained. Previous studies revealed that there are at least two classes of CO which occur in budding yeast and Arabidopsis <ref name="ref7"><ref name="ref8"><ref name="ref9">. Investigations of mer3 mutants in rice showed that apart from a reduced number of chiasmata, the remaining chiasmata distributed randomly among cells, suggesting that rice might also have at least two kinds of COs: one that appears to be sensitive to interference, whereas the other one is not <ref name="ref10">. The close number of residual chiasmata indicates similar contributions of HEI10 and MER3 during CO formation. Considering the close number and similar random distribution of residual chiasmata noted in mer3 and hei10 mutants, it is likely that HEI10 and MER3 may act in the same CO pathway, namely an interference-sensitive pathway.Besides a reduced chiasma frequency, hei10 also showed a precocious segregation of sister chromatids of some univalents at anaphase I.

===Labs working on this gene===
*State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
*Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

===References===
<references>
<ref name="ref1"> Toby GG, Gherraby W, Coleman TR, Golemis EA (2003) A novel RING finger protein, human enhancer of invasion 10, alters mitotic progression through regulation of cyclin B levels. Mol Cell Biol 23: 2109–2122. </ref>
<ref name="ref2"> Singh MK, Nicolas E, Gherraby W, Dadke D, Lessin S, et al. (2007) HEI10 negatively regulates cell invasion by inhibiting cyclin B/Cdk1 and other promotility proteins. Oncogene 26: 4825–4832. </ref>
<ref name="ref3">Wang, K., Wang, M., Tang, D., Shen, Y., Miao, C., Hu, Q., ... & Cheng, Z. (2012).The role of rice HEI10 in the formation of meiotic crossovers. PLoS genetics, 8(7), e1002809. </ref>
<ref name="ref4">de Boer E, Stam P, Dietrich AJ, Pastink A, Heyting C (2006) Two levels of interference in mouse meiotic recombination. Proc Natl Acad Sci U S A 103: 9607–9612. </ref>
<ref name="ref5">Lhuissier FG, Offenberg HH, Wittich PE, Vischer NO, Heyting C (2007) The mismatch repair protein MLH1 marks a subset of strongly interfering crossovers in tomato. Plant Cell 19: 862–876. </ref>
<ref name="ref6">Bhalla N, Wynne DJ, Jantsch V, Dernburg AF (2008) ZHP-3 acts at crossovers to couple meiotic recombination with synaptonemal complex disassembly and bivalent formation in C. elegans. PLoS Genet 4: e1000235. </ref>
<ref name="ref7">Borner GV, Kleckner N, Hunter N (2004) Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell 117: 29–45. </ref>
<ref name="ref8"> Mercier R, Jolivet S, Vezon D, Huppe E, Chelysheva L, et al. (2005) Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not. Curr Biol 15: 692–701. </ref>
<ref name="ref9">Higgins JD, Armstrong SJ, Franklin FC, Jones GH (2004) The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis. Genes & Dev 18: 2557–2570. </ref>
<ref name="ref10">Wang K, Tang D, Wang M, Lu J, Yu H, et al. (2009) MER3 is required for normal meiotic crossover formation, but not for presynaptic alignment in rice. J Cell Sci 122: 2055–2063. </ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

Os02g0232100

2014-06-04T10:19:45Z

Simon: /* Labs working on this gene */

HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.
===Mutation===
[[File:Figure111.jpg|right|thumb|150px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref3" />).'']]
60Co γ-ray irradiation was used to induce a sterile mutant from the japonica rice variety Wuxiangjing 9.The mutant showed normal vegetative growth but exhibited complete sterility (Figure 1A and 1B). Cytological observation of anthers showed that almost all pollens were shrunken and inviable (Figure 1C and 1D). When pollinated with wild-type (WT) pollens, the mutant spikelets were unable to set any seeds, suggesting that the female gametes were also sterile. The progenies of the heterozygous plants segregated from normal to sterile phenotype in a 3:1 ratio (fertile, 30; sterile, 10), indicating that a single recessive gene is responsible for the sterile phenotype<ref name="ref3" />.

===Expression（Mutant VS Wild type）===
[[File:Figure222.jpg|right|thumb|150px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure333.jpg|right|thumb|150px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure444.jpg|right|thumb|150px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.

===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.

===Labs working on this gene===
*State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
*Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

===References===
<references>
<ref name="ref1"> Toby GG, Gherraby W, Coleman TR, Golemis EA (2003) A novel RING finger protein, human enhancer of invasion 10, alters mitotic progression through regulation of cyclin B levels. Mol Cell Biol 23: 2109–2122. </ref>
<ref name="ref2"> Singh MK, Nicolas E, Gherraby W, Dadke D, Lessin S, et al. (2007) HEI10 negatively regulates cell invasion by inhibiting cyclin B/Cdk1 and other promotility proteins. Oncogene 26: 4825–4832. </ref>
<ref name="ref3">Wang, K., Wang, M., Tang, D., Shen, Y., Miao, C., Hu, Q., ... & Cheng, Z. (2012).The role of rice HEI10 in the formation of meiotic crossovers. PLoS genetics, 8(7), e1002809. </ref>
<ref name="ref4">de Boer E, Stam P, Dietrich AJ, Pastink A, Heyting C (2006) Two levels of interference in mouse meiotic recombination. Proc Natl Acad Sci U S A 103: 9607–9612. </ref>
<ref name="ref5">Lhuissier FG, Offenberg HH, Wittich PE, Vischer NO, Heyting C (2007) The mismatch repair protein MLH1 marks a subset of strongly interfering crossovers in tomato. Plant Cell 19: 862–876. </ref>
<ref name="ref6">Bhalla N, Wynne DJ, Jantsch V, Dernburg AF (2008) ZHP-3 acts at crossovers to couple meiotic recombination with synaptonemal complex disassembly and bivalent formation in C. elegans. PLoS Genet 4: e1000235. </ref>
<ref name="ref7">Borner GV, Kleckner N, Hunter N (2004) Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell 117: 29–45. </ref>
<ref name="ref8"> Mercier R, Jolivet S, Vezon D, Huppe E, Chelysheva L, et al. (2005) Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not. Curr Biol 15: 692–701. </ref>
<ref name="ref9">Higgins JD, Armstrong SJ, Franklin FC, Jones GH (2004) The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis. Genes & Dev 18: 2557–2570. </ref>
<ref name="ref10">Wang K, Tang D, Wang M, Lu J, Yu H, et al. (2009) MER3 is required for normal meiotic crossover formation, but not for presynaptic alignment in rice. J Cell Sci 122: 2055–2063. </ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

Os02g0232100

2014-06-04T10:19:11Z

Simon: /* Knowledge Extension */

HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.
===Mutation===
[[File:Figure111.jpg|right|thumb|150px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref3" />).'']]
60Co γ-ray irradiation was used to induce a sterile mutant from the japonica rice variety Wuxiangjing 9.The mutant showed normal vegetative growth but exhibited complete sterility (Figure 1A and 1B). Cytological observation of anthers showed that almost all pollens were shrunken and inviable (Figure 1C and 1D). When pollinated with wild-type (WT) pollens, the mutant spikelets were unable to set any seeds, suggesting that the female gametes were also sterile. The progenies of the heterozygous plants segregated from normal to sterile phenotype in a 3:1 ratio (fertile, 30; sterile, 10), indicating that a single recessive gene is responsible for the sterile phenotype<ref name="ref3" />.

===Expression（Mutant VS Wild type）===
[[File:Figure222.jpg|right|thumb|150px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure333.jpg|right|thumb|150px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure444.jpg|right|thumb|150px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.

===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.

===Labs working on this gene===
State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

===References===
<references>
<ref name="ref1"> Toby GG, Gherraby W, Coleman TR, Golemis EA (2003) A novel RING finger protein, human enhancer of invasion 10, alters mitotic progression through regulation of cyclin B levels. Mol Cell Biol 23: 2109–2122. </ref>
<ref name="ref2"> Singh MK, Nicolas E, Gherraby W, Dadke D, Lessin S, et al. (2007) HEI10 negatively regulates cell invasion by inhibiting cyclin B/Cdk1 and other promotility proteins. Oncogene 26: 4825–4832. </ref>
<ref name="ref3">Wang, K., Wang, M., Tang, D., Shen, Y., Miao, C., Hu, Q., ... & Cheng, Z. (2012).The role of rice HEI10 in the formation of meiotic crossovers. PLoS genetics, 8(7), e1002809. </ref>
<ref name="ref4">de Boer E, Stam P, Dietrich AJ, Pastink A, Heyting C (2006) Two levels of interference in mouse meiotic recombination. Proc Natl Acad Sci U S A 103: 9607–9612. </ref>
<ref name="ref5">Lhuissier FG, Offenberg HH, Wittich PE, Vischer NO, Heyting C (2007) The mismatch repair protein MLH1 marks a subset of strongly interfering crossovers in tomato. Plant Cell 19: 862–876. </ref>
<ref name="ref6">Bhalla N, Wynne DJ, Jantsch V, Dernburg AF (2008) ZHP-3 acts at crossovers to couple meiotic recombination with synaptonemal complex disassembly and bivalent formation in C. elegans. PLoS Genet 4: e1000235. </ref>
<ref name="ref7">Borner GV, Kleckner N, Hunter N (2004) Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell 117: 29–45. </ref>
<ref name="ref8"> Mercier R, Jolivet S, Vezon D, Huppe E, Chelysheva L, et al. (2005) Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not. Curr Biol 15: 692–701. </ref>
<ref name="ref9">Higgins JD, Armstrong SJ, Franklin FC, Jones GH (2004) The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis. Genes & Dev 18: 2557–2570. </ref>
<ref name="ref10">Wang K, Tang D, Wang M, Lu J, Yu H, et al. (2009) MER3 is required for normal meiotic crossover formation, but not for presynaptic alignment in rice. J Cell Sci 122: 2055–2063. </ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

Os02g0232100

2014-06-04T10:16:14Z

Simon:

HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.
===Mutation===
[[File:Figure111.jpg|right|thumb|150px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref3" />).'']]
60Co γ-ray irradiation was used to induce a sterile mutant from the japonica rice variety Wuxiangjing 9.The mutant showed normal vegetative growth but exhibited complete sterility (Figure 1A and 1B). Cytological observation of anthers showed that almost all pollens were shrunken and inviable (Figure 1C and 1D). When pollinated with wild-type (WT) pollens, the mutant spikelets were unable to set any seeds, suggesting that the female gametes were also sterile. The progenies of the heterozygous plants segregated from normal to sterile phenotype in a 3:1 ratio (fertile, 30; sterile, 10), indicating that a single recessive gene is responsible for the sterile phenotype<ref name="ref3" />.

===Expression（Mutant VS Wild type）===
[[File:Figure222.jpg|right|thumb|150px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure333.jpg|right|thumb|150px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure444.jpg|right|thumb|150px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.

===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.

===Knowledge Extension===
In hei10 mutants, only about 31% chiasmata were maintained. This number is similar to that of mer3 mutants, in which about 28% chiasmata were remained. Previous studies revealed that there are at least two classes of CO which occur in budding yeast and Arabidopsis <ref name="ref7"><ref name="ref8"><ref name="ref9">. Investigations of mer3 mutants in rice showed that apart from a reduced number of chiasmata, the remaining chiasmata distributed randomly among cells, suggesting that rice might also have at least two kinds of COs: one that appears to be sensitive to interference, whereas the other one is not <ref name="ref10">. The close number of residual chiasmata indicates similar contributions of HEI10 and MER3 during CO formation. Considering the close number and similar random distribution of residual chiasmata noted in mer3 and hei10 mutants, it is likely that HEI10 and MER3 may act in the same CO pathway, namely an interference-sensitive pathway.Besides a reduced chiasma frequency, hei10 also showed a precocious segregation of sister chromatids of some univalents at anaphase I.

===Labs working on this gene===
State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

===References===
<references>
<ref name="ref1"> Toby GG, Gherraby W, Coleman TR, Golemis EA (2003) A novel RING finger protein, human enhancer of invasion 10, alters mitotic progression through regulation of cyclin B levels. Mol Cell Biol 23: 2109–2122. </ref>
<ref name="ref2"> Singh MK, Nicolas E, Gherraby W, Dadke D, Lessin S, et al. (2007) HEI10 negatively regulates cell invasion by inhibiting cyclin B/Cdk1 and other promotility proteins. Oncogene 26: 4825–4832. </ref>
<ref name="ref3">Wang, K., Wang, M., Tang, D., Shen, Y., Miao, C., Hu, Q., ... & Cheng, Z. (2012).The role of rice HEI10 in the formation of meiotic crossovers. PLoS genetics, 8(7), e1002809. </ref>
<ref name="ref4">de Boer E, Stam P, Dietrich AJ, Pastink A, Heyting C (2006) Two levels of interference in mouse meiotic recombination. Proc Natl Acad Sci U S A 103: 9607–9612. </ref>
<ref name="ref5">Lhuissier FG, Offenberg HH, Wittich PE, Vischer NO, Heyting C (2007) The mismatch repair protein MLH1 marks a subset of strongly interfering crossovers in tomato. Plant Cell 19: 862–876. </ref>
<ref name="ref6">Bhalla N, Wynne DJ, Jantsch V, Dernburg AF (2008) ZHP-3 acts at crossovers to couple meiotic recombination with synaptonemal complex disassembly and bivalent formation in C. elegans. PLoS Genet 4: e1000235. </ref>
<ref name="ref7">Borner GV, Kleckner N, Hunter N (2004) Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell 117: 29–45. </ref>
<ref name="ref8"> Mercier R, Jolivet S, Vezon D, Huppe E, Chelysheva L, et al. (2005) Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not. Curr Biol 15: 692–701. </ref>
<ref name="ref9">Higgins JD, Armstrong SJ, Franklin FC, Jones GH (2004) The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis. Genes & Dev 18: 2557–2570. </ref>
<ref name="ref10">Wang K, Tang D, Wang M, Lu J, Yu H, et al. (2009) MER3 is required for normal meiotic crossover formation, but not for presynaptic alignment in rice. J Cell Sci 122: 2055–2063. </ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

Os02g0232100

2014-06-04T10:14:44Z

Simon: /* Annotated Information */

Please input one-sentence summary here.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.
===Mutation===
[[File:Figure111.jpg|right|thumb|150px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref3" />).'']]
60Co γ-ray irradiation was used to induce a sterile mutant from the japonica rice variety Wuxiangjing 9.The mutant showed normal vegetative growth but exhibited complete sterility (Figure 1A and 1B). Cytological observation of anthers showed that almost all pollens were shrunken and inviable (Figure 1C and 1D). When pollinated with wild-type (WT) pollens, the mutant spikelets were unable to set any seeds, suggesting that the female gametes were also sterile. The progenies of the heterozygous plants segregated from normal to sterile phenotype in a 3:1 ratio (fertile, 30; sterile, 10), indicating that a single recessive gene is responsible for the sterile phenotype<ref name="ref3" />.

===Expression（Mutant VS Wild type）===
[[File:Figure222.jpg|right|thumb|150px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure333.jpg|right|thumb|150px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure444.jpg|right|thumb|150px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.

===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.
===Knowledge Extension===

In hei10 mutants, only about 31% chiasmata were maintained. This number is similar to that of mer3 mutants, in which about 28% chiasmata were remained. Previous studies revealed that there are at least two classes of CO which occur in budding yeast and Arabidopsis <ref name="ref7"><ref name="ref8"><ref name="ref9">. Investigations of mer3 mutants in rice showed that apart from a reduced number of chiasmata, the remaining chiasmata distributed randomly among cells, suggesting that rice might also have at least two kinds of COs: one that appears to be sensitive to interference, whereas the other one is not <ref name="ref10">. The close number of residual chiasmata indicates similar contributions of HEI10 and MER3 during CO formation. Considering the close number and similar random distribution of residual chiasmata noted in mer3 and hei10 mutants, it is likely that HEI10 and MER3 may act in the same CO pathway, namely an interference-sensitive pathway.Besides a reduced chiasma frequency, hei10 also showed a precocious segregation of sister chromatids of some univalents at anaphase I.

===Labs working on this gene===
State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

===References===
<references>
<ref name="ref1"> Toby GG, Gherraby W, Coleman TR, Golemis EA (2003) A novel RING finger protein, human enhancer of invasion 10, alters mitotic progression through regulation of cyclin B levels. Mol Cell Biol 23: 2109–2122. </ref>
<ref name="ref2"> Singh MK, Nicolas E, Gherraby W, Dadke D, Lessin S, et al. (2007) HEI10 negatively regulates cell invasion by inhibiting cyclin B/Cdk1 and other promotility proteins. Oncogene 26: 4825–4832. </ref>
<ref name="ref3">Wang, K., Wang, M., Tang, D., Shen, Y., Miao, C., Hu, Q., ... & Cheng, Z. (2012).The role of rice HEI10 in the formation of meiotic crossovers. PLoS genetics, 8(7), e1002809. </ref>
<ref name="ref4">de Boer E, Stam P, Dietrich AJ, Pastink A, Heyting C (2006) Two levels of interference in mouse meiotic recombination. Proc Natl Acad Sci U S A 103: 9607–9612. </ref>
<ref name="ref5">Lhuissier FG, Offenberg HH, Wittich PE, Vischer NO, Heyting C (2007) The mismatch repair protein MLH1 marks a subset of strongly interfering crossovers in tomato. Plant Cell 19: 862–876. </ref>
<ref name="ref6">Bhalla N, Wynne DJ, Jantsch V, Dernburg AF (2008) ZHP-3 acts at crossovers to couple meiotic recombination with synaptonemal complex disassembly and bivalent formation in C. elegans. PLoS Genet 4: e1000235. </ref>
<ref name="ref7">Borner GV, Kleckner N, Hunter N (2004) Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell 117: 29–45. </ref>
<ref name="ref8"> Mercier R, Jolivet S, Vezon D, Huppe E, Chelysheva L, et al. (2005) Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not. Curr Biol 15: 692–701. </ref>
<ref name="ref9">Higgins JD, Armstrong SJ, Franklin FC, Jones GH (2004) The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis. Genes & Dev 18: 2557–2570. </ref>
<ref name="ref10">Wang K, Tang D, Wang M, Lu J, Yu H, et al. (2009) MER3 is required for normal meiotic crossover formation, but not for presynaptic alignment in rice. J Cell Sci 122: 2055–2063. </ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

Os02g0232100

2014-06-04T10:13:52Z

Simon: /* Annotated Information */

Please input one-sentence summary here.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.
===Mutation===
[[File:Figure111.jpg|right|thumb|150px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref3" />).'']]
60Co γ-ray irradiation was used to induce a sterile mutant from the japonica rice variety Wuxiangjing 9.The mutant showed normal vegetative growth but exhibited complete sterility (Figure 1A and 1B). Cytological observation of anthers showed that almost all pollens were shrunken and inviable (Figure 1C and 1D). When pollinated with wild-type (WT) pollens, the mutant spikelets were unable to set any seeds, suggesting that the female gametes were also sterile. The progenies of the heterozygous plants segregated from normal to sterile phenotype in a 3:1 ratio (fertile, 30; sterile, 10), indicating that a single recessive gene is responsible for the sterile phenotype<ref name="ref3" />.

===Expression（Mutant VS Wild type）===
[[File:Figure222.jpg|right|thumb|150px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure333.jpg|right|thumb|150px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure444.jpg|right|thumb|150px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.

===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.
===Knowledge Extension===

In hei10 mutants, only about 31% chiasmata were maintained. This number is similar to that of mer3 mutants, in which about 28% chiasmata were remained. Previous studies revealed that there are at least two classes of CO which occur in budding yeast and Arabidopsis <ref name="ref7"><ref name="ref8"><ref name="ref9">. Investigations of mer3 mutants in rice showed that apart from a reduced number of chiasmata, the remaining chiasmata distributed randomly among cells, suggesting that rice might also have at least two kinds of COs: one that appears to be sensitive to interference, whereas the other one is not <ref name="ref10">. The close number of residual chiasmata indicates similar contributions of HEI10 and MER3 during CO formation. Considering the close number and similar random distribution of residual chiasmata noted in mer3 and hei10 mutants, it is likely that HEI10 and MER3 may act in the same CO pathway, namely an interference-sensitive pathway.Besides a reduced chiasma frequency, hei10 also showed a precocious segregation of sister chromatids of some univalents at anaphase I.

==Labs working on this gene==
State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

==References==
<references>
<ref name="ref1"> Toby GG, Gherraby W, Coleman TR, Golemis EA (2003) A novel RING finger protein, human enhancer of invasion 10, alters mitotic progression through regulation of cyclin B levels. Mol Cell Biol 23: 2109–2122. </ref>
<ref name="ref2"> Singh MK, Nicolas E, Gherraby W, Dadke D, Lessin S, et al. (2007) HEI10 negatively regulates cell invasion by inhibiting cyclin B/Cdk1 and other promotility proteins. Oncogene 26: 4825–4832. </ref>
<ref name="ref3">Wang, K., Wang, M., Tang, D., Shen, Y., Miao, C., Hu, Q., ... & Cheng, Z. (2012).The role of rice HEI10 in the formation of meiotic crossovers. PLoS genetics, 8(7), e1002809. </ref>
<ref name="ref4">de Boer E, Stam P, Dietrich AJ, Pastink A, Heyting C (2006) Two levels of interference in mouse meiotic recombination. Proc Natl Acad Sci U S A 103: 9607–9612. </ref>
<ref name="ref5">Lhuissier FG, Offenberg HH, Wittich PE, Vischer NO, Heyting C (2007) The mismatch repair protein MLH1 marks a subset of strongly interfering crossovers in tomato. Plant Cell 19: 862–876. </ref>
<ref name="ref6">Bhalla N, Wynne DJ, Jantsch V, Dernburg AF (2008) ZHP-3 acts at crossovers to couple meiotic recombination with synaptonemal complex disassembly and bivalent formation in C. elegans. PLoS Genet 4: e1000235. </ref>
<ref name="ref7">Borner GV, Kleckner N, Hunter N (2004) Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell 117: 29–45. </ref>
<ref name="ref8"> Mercier R, Jolivet S, Vezon D, Huppe E, Chelysheva L, et al. (2005) Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not. Curr Biol 15: 692–701. </ref>
<ref name="ref9">Higgins JD, Armstrong SJ, Franklin FC, Jones GH (2004) The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis. Genes & Dev 18: 2557–2570. </ref>
<ref name="ref10">Wang K, Tang D, Wang M, Lu J, Yu H, et al. (2009) MER3 is required for normal meiotic crossover formation, but not for presynaptic alignment in rice. J Cell Sci 122: 2055–2063. </ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

File:Figure444.jpg

2014-06-04T10:12:30Z

Simon: (A) Dual immunolocalization of WT scattered chromosomes with REC8 (red) and HEI10 (green) antibodies. Scale bar: 5 mm. (B) Dual immunolocalization of CENH3 and HEI10 at late pachytene. (C) No HEI10 signals were detected at metaphase I, chromosomes are sta

(A) Dual immunolocalization of WT scattered chromosomes with REC8 (red) and HEI10 (green) antibodies. Scale bar: 5 mm. (B) Dual immunolocalization of CENH3 and HEI10 at late pachytene. (C) No HEI10 signals were detected at metaphase I, chromosomes are stained with DAPI (blue). Scale bars: 5 mm. (D) Histogram of the observed distances between two adjacent HEI10 bright foci on the shortest chromosomes (percentages of the length of the synaptonemal complex). The gray bars show the observed relative frequencies of inter-HEI10 foci distances. The red curve presents the best fit of the distance to the gamma distribution. n is the interference parameter in the gamma model. v ̂ is the n value (estimated SE) for which the best fit of the observed distances to the gamma model was got. Here, the estimatev ̂ is 8.39, which indicates a strong interference among HEI10 bright foci on the shortest chromosome in rice (if there is no interference, n is 1).

File:Figure333.jpg

2014-06-04T10:11:33Z

Simon: (A) Pachytene. (B) Diakinesis with both bivalents and univalents. (C) Metaphase I with five bivalents and fourteen univalents. (D) Metaphase I with four bivalents and sixteen univalents. Thirteen univalents and all bivalents align on the equatorial plate.

(A) Pachytene. (B) Diakinesis with both bivalents and univalents. (C) Metaphase I with five bivalents and fourteen univalents. (D) Metaphase I with four bivalents and sixteen univalents. Thirteen univalents and all bivalents align on the equatorial plate. (E) Anaphase I. Many univalents undergo a precocious segregation of sister chromatids. (F) Telophase II. Tetrads with several randomly distributed chromosomes. Chromosomes were stained with DAPI. Scale bars, 5 mm.

File:Figure222.jpg

2014-06-04T10:10:52Z

Simon: uploaded a new version of "File:Figure222.jpg": (A) Schematic representation of HEI10 gene and the location of hei10 mutation. Translated regions are represented by black boxes and untranslated regions are indicated by white boxes. HEI10A an

(A) Schematic representation of HEI10 gene and the location of hei10 mutation. Translated regions are represented by black boxes and untranslated regions are indicated by white boxes. HEI10A and HEI10B indicate long and short transcripts, respectively.

File:Figure222.jpg

2014-06-04T10:10:51Z

Simon: (A) Schematic representation of HEI10 gene and the location of hei10 mutation. Translated regions are represented by black boxes and untranslated regions are indicated by white boxes. HEI10A and HEI10B indicate long and short transcripts, respectively.

File:Figure111.jpg

2014-06-04T10:10:08Z

Simon: (A) Comparison of a WT plant (left) and a hei10-1 mutant plant (right). (B) Comparison of a WT panicle (left) and a hei10-1 mutant panicle (right). (C) Fertile pollen grains in a WT plant. (D) Sterile pollen grains in a hei10-1 mutant plant. Scale bars: 5

(A) Comparison of a WT plant (left) and a hei10-1 mutant plant (right). (B) Comparison of a WT panicle (left) and a hei10-1 mutant panicle (right). (C) Fertile pollen grains in a WT plant. (D) Sterile pollen grains in a hei10-1 mutant plant. Scale bars: 50 µm.

Os02g0232100

2014-06-04T10:09:11Z

Simon: /* Annotated Information */

Please input one-sentence summary here.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.
===Mutation===
[[File:Figure111.jpg|right|thumb|150px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref3" />).'']]
60Co γ-ray irradiation was used to induce a sterile mutant from the japonica rice variety Wuxiangjing 9.The mutant showed normal vegetative growth but exhibited complete sterility (Figure 1A and 1B). Cytological observation of anthers showed that almost all pollens were shrunken and inviable (Figure 1C and 1D). When pollinated with wild-type (WT) pollens, the mutant spikelets were unable to set any seeds, suggesting that the female gametes were also sterile. The progenies of the heterozygous plants segregated from normal to sterile phenotype in a 3:1 ratio (fertile, 30; sterile, 10), indicating that a single recessive gene is responsible for the sterile phenotype<ref name="ref3" />.

===Expression（Mutant VS Wild type）===
[[File:Figure222.jpg|right|thumb|150px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure333.jpg|right|thumb|150px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure444.jpg|right|thumb|150px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.

===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.

==Labs working on this gene==
State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

==References==
<references>
<ref name="ref1"> Toby GG, Gherraby W, Coleman TR, Golemis EA (2003) A novel RING finger protein, human enhancer of invasion 10, alters mitotic progression through regulation of cyclin B levels. Mol Cell Biol 23: 2109–2122. </ref>
<ref name="ref2"> Singh MK, Nicolas E, Gherraby W, Dadke D, Lessin S, et al. (2007) HEI10 negatively regulates cell invasion by inhibiting cyclin B/Cdk1 and other promotility proteins. Oncogene 26: 4825–4832. </ref>
<ref name="ref3">Wang, K., Wang, M., Tang, D., Shen, Y., Miao, C., Hu, Q., ... & Cheng, Z. (2012).The role of rice HEI10 in the formation of meiotic crossovers. PLoS genetics, 8(7), e1002809. </ref>
<ref name="ref4">de Boer E, Stam P, Dietrich AJ, Pastink A, Heyting C (2006) Two levels of interference in mouse meiotic recombination. Proc Natl Acad Sci U S A 103: 9607–9612. </ref>
<ref name="ref5">Lhuissier FG, Offenberg HH, Wittich PE, Vischer NO, Heyting C (2007) The mismatch repair protein MLH1 marks a subset of strongly interfering crossovers in tomato. Plant Cell 19: 862–876. </ref>
<ref name="ref6">Bhalla N, Wynne DJ, Jantsch V, Dernburg AF (2008) ZHP-3 acts at crossovers to couple meiotic recombination with synaptonemal complex disassembly and bivalent formation in C. elegans. PLoS Genet 4: e1000235. </ref>
<ref name="ref7">Borner GV, Kleckner N, Hunter N (2004) Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell 117: 29–45. </ref>
<ref name="ref8"> Mercier R, Jolivet S, Vezon D, Huppe E, Chelysheva L, et al. (2005) Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not. Curr Biol 15: 692–701. </ref>
<ref name="ref9">Higgins JD, Armstrong SJ, Franklin FC, Jones GH (2004) The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis. Genes & Dev 18: 2557–2570. </ref>
<ref name="ref10">Wang K, Tang D, Wang M, Lu J, Yu H, et al. (2009) MER3 is required for normal meiotic crossover formation, but not for presynaptic alignment in rice. J Cell Sci 122: 2055–2063. </ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

Os02g0232100

2014-06-04T10:08:43Z

Simon: /* Annotated Information */

Please input one-sentence summary here.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.
===Mutation===
[[File:Figure1111.jpg|right|thumb|150px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref3" />).'']]
60Co γ-ray irradiation was used to induce a sterile mutant from the japonica rice variety Wuxiangjing 9.The mutant showed normal vegetative growth but exhibited complete sterility (Figure 1A and 1B). Cytological observation of anthers showed that almost all pollens were shrunken and inviable (Figure 1C and 1D). When pollinated with wild-type (WT) pollens, the mutant spikelets were unable to set any seeds, suggesting that the female gametes were also sterile. The progenies of the heterozygous plants segregated from normal to sterile phenotype in a 3:1 ratio (fertile, 30; sterile, 10), indicating that a single recessive gene is responsible for the sterile phenotype<ref name="ref3" />.

===Expression（Mutant VS Wild type）===
[[File:Figure2222.jpg|right|thumb|150px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure3333.jpg|right|thumb|150px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure4444.jpg|right|thumb|150px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.

===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.

==Labs working on this gene==
State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

==References==
<references>
<ref name="ref1"> Toby GG, Gherraby W, Coleman TR, Golemis EA (2003) A novel RING finger protein, human enhancer of invasion 10, alters mitotic progression through regulation of cyclin B levels. Mol Cell Biol 23: 2109–2122. </ref>
<ref name="ref2"> Singh MK, Nicolas E, Gherraby W, Dadke D, Lessin S, et al. (2007) HEI10 negatively regulates cell invasion by inhibiting cyclin B/Cdk1 and other promotility proteins. Oncogene 26: 4825–4832. </ref>
<ref name="ref3">Wang, K., Wang, M., Tang, D., Shen, Y., Miao, C., Hu, Q., ... & Cheng, Z. (2012).The role of rice HEI10 in the formation of meiotic crossovers. PLoS genetics, 8(7), e1002809. </ref>
<ref name="ref4">de Boer E, Stam P, Dietrich AJ, Pastink A, Heyting C (2006) Two levels of interference in mouse meiotic recombination. Proc Natl Acad Sci U S A 103: 9607–9612. </ref>
<ref name="ref5">Lhuissier FG, Offenberg HH, Wittich PE, Vischer NO, Heyting C (2007) The mismatch repair protein MLH1 marks a subset of strongly interfering crossovers in tomato. Plant Cell 19: 862–876. </ref>
<ref name="ref6">Bhalla N, Wynne DJ, Jantsch V, Dernburg AF (2008) ZHP-3 acts at crossovers to couple meiotic recombination with synaptonemal complex disassembly and bivalent formation in C. elegans. PLoS Genet 4: e1000235. </ref>
<ref name="ref7">Borner GV, Kleckner N, Hunter N (2004) Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell 117: 29–45. </ref>
<ref name="ref8"> Mercier R, Jolivet S, Vezon D, Huppe E, Chelysheva L, et al. (2005) Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not. Curr Biol 15: 692–701. </ref>
<ref name="ref9">Higgins JD, Armstrong SJ, Franklin FC, Jones GH (2004) The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis. Genes & Dev 18: 2557–2570. </ref>
<ref name="ref10">Wang K, Tang D, Wang M, Lu J, Yu H, et al. (2009) MER3 is required for normal meiotic crossover formation, but not for presynaptic alignment in rice. J Cell Sci 122: 2055–2063. </ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

Os02g0232100

2014-06-04T10:04:14Z

Simon: /* Expression（Mutant VS Wild type） */

Please input one-sentence summary here.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.
===Mutation===
[[File:Figure1.jpg|right|thumb|250px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref3" />).'']]
60Co γ-ray irradiation was used to induce a sterile mutant from the japonica rice variety Wuxiangjing 9.The mutant showed normal vegetative growth but exhibited complete sterility (Figure 1A and 1B). Cytological observation of anthers showed that almost all pollens were shrunken and inviable (Figure 1C and 1D). When pollinated with wild-type (WT) pollens, the mutant spikelets were unable to set any seeds, suggesting that the female gametes were also sterile. The progenies of the heterozygous plants segregated from normal to sterile phenotype in a 3:1 ratio (fertile, 30; sterile, 10), indicating that a single recessive gene is responsible for the sterile phenotype<ref name="ref3" />.

===Expression（Mutant VS Wild type）===
[[File:Figure2.jpg|right|thumb|250px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure3.jpg|right|thumb|250px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure4.jpg|right|thumb|250px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.

===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.

==Labs working on this gene==
State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

==References==
<references>
<ref name="ref1"> Toby GG, Gherraby W, Coleman TR, Golemis EA (2003) A novel RING finger protein, human enhancer of invasion 10, alters mitotic progression through regulation of cyclin B levels. Mol Cell Biol 23: 2109–2122. </ref>
<ref name="ref2"> Singh MK, Nicolas E, Gherraby W, Dadke D, Lessin S, et al. (2007) HEI10 negatively regulates cell invasion by inhibiting cyclin B/Cdk1 and other promotility proteins. Oncogene 26: 4825–4832. </ref>
<ref name="ref3">Wang, K., Wang, M., Tang, D., Shen, Y., Miao, C., Hu, Q., ... & Cheng, Z. (2012).The role of rice HEI10 in the formation of meiotic crossovers. PLoS genetics, 8(7), e1002809. </ref>
<ref name="ref4">de Boer E, Stam P, Dietrich AJ, Pastink A, Heyting C (2006) Two levels of interference in mouse meiotic recombination. Proc Natl Acad Sci U S A 103: 9607–9612. </ref>
<ref name="ref5">Lhuissier FG, Offenberg HH, Wittich PE, Vischer NO, Heyting C (2007) The mismatch repair protein MLH1 marks a subset of strongly interfering crossovers in tomato. Plant Cell 19: 862–876. </ref>
<ref name="ref6">Bhalla N, Wynne DJ, Jantsch V, Dernburg AF (2008) ZHP-3 acts at crossovers to couple meiotic recombination with synaptonemal complex disassembly and bivalent formation in C. elegans. PLoS Genet 4: e1000235. </ref>
<ref name="ref7">Borner GV, Kleckner N, Hunter N (2004) Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell 117: 29–45. </ref>
<ref name="ref8"> Mercier R, Jolivet S, Vezon D, Huppe E, Chelysheva L, et al. (2005) Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not. Curr Biol 15: 692–701. </ref>
<ref name="ref9">Higgins JD, Armstrong SJ, Franklin FC, Jones GH (2004) The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis. Genes & Dev 18: 2557–2570. </ref>
<ref name="ref10">Wang K, Tang D, Wang M, Lu J, Yu H, et al. (2009) MER3 is required for normal meiotic crossover formation, but not for presynaptic alignment in rice. J Cell Sci 122: 2055–2063. </ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

File:Figure 4. Analysis of the distribution of HEI10 bright foci in WT meiocytes.jpg

2014-06-04T10:02:11Z

File:Figure 3.Meiosis in the hei10-1 mutant.jpg

2014-06-04T10:00:50Z

Os02g0232100

2014-06-04T09:59:35Z

Simon: /* Mutation */

Please input one-sentence summary here.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.
===Mutation===
[[File:Figure1.jpg|right|thumb|250px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref3" />).'']]
60Co γ-ray irradiation was used to induce a sterile mutant from the japonica rice variety Wuxiangjing 9.The mutant showed normal vegetative growth but exhibited complete sterility (Figure 1A and 1B). Cytological observation of anthers showed that almost all pollens were shrunken and inviable (Figure 1C and 1D). When pollinated with wild-type (WT) pollens, the mutant spikelets were unable to set any seeds, suggesting that the female gametes were also sterile. The progenies of the heterozygous plants segregated from normal to sterile phenotype in a 3:1 ratio (fertile, 30; sterile, 10), indicating that a single recessive gene is responsible for the sterile phenotype<ref name="ref3" />.

===Expression（Mutant VS Wild type）===
[[File:Figure222.jpg|right|thumb|250px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure333.jpg|right|thumb|250px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure444.jpg|right|thumb|250px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.

===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.

==Labs working on this gene==
State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

==References==
<references>
<ref name="ref1"> Toby GG, Gherraby W, Coleman TR, Golemis EA (2003) A novel RING finger protein, human enhancer of invasion 10, alters mitotic progression through regulation of cyclin B levels. Mol Cell Biol 23: 2109–2122. </ref>
<ref name="ref2"> Singh MK, Nicolas E, Gherraby W, Dadke D, Lessin S, et al. (2007) HEI10 negatively regulates cell invasion by inhibiting cyclin B/Cdk1 and other promotility proteins. Oncogene 26: 4825–4832. </ref>
<ref name="ref3">Wang, K., Wang, M., Tang, D., Shen, Y., Miao, C., Hu, Q., ... & Cheng, Z. (2012).The role of rice HEI10 in the formation of meiotic crossovers. PLoS genetics, 8(7), e1002809. </ref>
<ref name="ref4">de Boer E, Stam P, Dietrich AJ, Pastink A, Heyting C (2006) Two levels of interference in mouse meiotic recombination. Proc Natl Acad Sci U S A 103: 9607–9612. </ref>
<ref name="ref5">Lhuissier FG, Offenberg HH, Wittich PE, Vischer NO, Heyting C (2007) The mismatch repair protein MLH1 marks a subset of strongly interfering crossovers in tomato. Plant Cell 19: 862–876. </ref>
<ref name="ref6">Bhalla N, Wynne DJ, Jantsch V, Dernburg AF (2008) ZHP-3 acts at crossovers to couple meiotic recombination with synaptonemal complex disassembly and bivalent formation in C. elegans. PLoS Genet 4: e1000235. </ref>
<ref name="ref7">Borner GV, Kleckner N, Hunter N (2004) Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell 117: 29–45. </ref>
<ref name="ref8"> Mercier R, Jolivet S, Vezon D, Huppe E, Chelysheva L, et al. (2005) Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not. Curr Biol 15: 692–701. </ref>
<ref name="ref9">Higgins JD, Armstrong SJ, Franklin FC, Jones GH (2004) The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis. Genes & Dev 18: 2557–2570. </ref>
<ref name="ref10">Wang K, Tang D, Wang M, Lu J, Yu H, et al. (2009) MER3 is required for normal meiotic crossover formation, but not for presynaptic alignment in rice. J Cell Sci 122: 2055–2063. </ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

File:Figure 2.Organization of the HEI10 gene and protein alignment.jpg

2014-06-04T09:52:42Z

Simon:

[[File:Figure2.jpg|right|thumb|250px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']] (A) Schematic representation of HEI10 gene and the location of hei10 mutation. Translated regions are represented by black boxes and untranslated regions are indicated by white boxes. HEI10A and HEI10B indicate long and short transcripts, respectively.
doi:10.1371/journal.pgen.1002809.g001

File:Figure 2.Organization of the HEI10 gene and protein alignment.jpg

2014-06-04T09:51:51Z

File:The rice hei10-1 mutant phenotype.jpg

2014-06-04T09:50:01Z

Simon: uploaded a new version of "File:The rice hei10-1 mutant phenotype.jpg": (A) Comparison of a WT plant (left) and a hei10-1 mutant plant (right). (B) Comparison of a WT panicle (left) and a hei10-1 mutant panicle (right). (C) Fertile pollen gr

[[File:Figure1.jpg|right|thumb|250px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref1" />).'']](A) Comparison of a WT plant (left) and a hei10-1 mutant plant (right). (B) Comparison of a WT panicle (left) and a hei10-1 mutant panicle (right). (C) Fertile pollen grains in a WT plant. (D) Sterile pollen grains in a hei10-1 mutant plant. Scale bars: 50 µm.
doi:10.1371/journal.pgen.1002809.s001

File:The rice hei10-1 mutant phenotype.jpg

2014-06-04T09:37:38Z

Simon:

File:The rice hei10-1 mutant phenotype.jpg

2014-06-04T09:34:52Z

Simon:

[[File:Figure1.jpg|right|thumb|250px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref3" />).'']](A) Comparison of a WT plant (left) and a hei10-1 mutant plant (right). (B) Comparison of a WT panicle (left) and a hei10-1 mutant panicle (right). (C) Fertile pollen grains in a WT plant. (D) Sterile pollen grains in a hei10-1 mutant plant. Scale bars: 50 µm.
doi:10.1371/journal.pgen.1002809.s001

File:The rice hei10-1 mutant phenotype.jpg

2014-06-04T09:32:31Z

Simon:

[[File:The rice hei10-1 mutant phenotype.jpg]](A) Comparison of a WT plant (left) and a hei10-1 mutant plant (right). (B) Comparison of a WT panicle (left) and a hei10-1 mutant panicle (right). (C) Fertile pollen grains in a WT plant. (D) Sterile pollen grains in a hei10-1 mutant plant. Scale bars: 50 µm.
doi:10.1371/journal.pgen.1002809.s001

File:The rice hei10-1 mutant phenotype.jpg

2014-06-04T09:31:22Z

File:Figure 1.The rice hei10-1 mutant phenotype.jpg

2014-06-04T09:18:17Z

Simon:

File:Figure 1.The rice hei10-1 mutant phenotype.jpg

2014-06-04T09:14:41Z

Os02g0232100

2014-06-04T09:07:39Z

Simon: /* Expression（Mutant VS Wild type） */

Please input one-sentence summary here.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.
===Mutation===
[[File:Figure111.jpg|right|thumb|250px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref3" />).'']]
60Co γ-ray irradiation was used to induce a sterile mutant from the japonica rice variety Wuxiangjing 9.The mutant showed normal vegetative growth but exhibited complete sterility (Figure 1A and 1B). Cytological observation of anthers showed that almost all pollens were shrunken and inviable (Figure 1C and 1D). When pollinated with wild-type (WT) pollens, the mutant spikelets were unable to set any seeds, suggesting that the female gametes were also sterile. The progenies of the heterozygous plants segregated from normal to sterile phenotype in a 3:1 ratio (fertile, 30; sterile, 10), indicating that a single recessive gene is responsible for the sterile phenotype<ref name="ref3" />.

===Expression（Mutant VS Wild type）===
[[File:Figure222.jpg|right|thumb|250px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure333.jpg|right|thumb|250px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure444.jpg|right|thumb|250px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.

===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.

==Labs working on this gene==
State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

==References==
<references>
<ref name="ref1"> Toby GG, Gherraby W, Coleman TR, Golemis EA (2003) A novel RING finger protein, human enhancer of invasion 10, alters mitotic progression through regulation of cyclin B levels. Mol Cell Biol 23: 2109–2122. </ref>
<ref name="ref2"> Singh MK, Nicolas E, Gherraby W, Dadke D, Lessin S, et al. (2007) HEI10 negatively regulates cell invasion by inhibiting cyclin B/Cdk1 and other promotility proteins. Oncogene 26: 4825–4832. </ref>
<ref name="ref3">Wang, K., Wang, M., Tang, D., Shen, Y., Miao, C., Hu, Q., ... & Cheng, Z. (2012).The role of rice HEI10 in the formation of meiotic crossovers. PLoS genetics, 8(7), e1002809. </ref>
<ref name="ref4">de Boer E, Stam P, Dietrich AJ, Pastink A, Heyting C (2006) Two levels of interference in mouse meiotic recombination. Proc Natl Acad Sci U S A 103: 9607–9612. </ref>
<ref name="ref5">Lhuissier FG, Offenberg HH, Wittich PE, Vischer NO, Heyting C (2007) The mismatch repair protein MLH1 marks a subset of strongly interfering crossovers in tomato. Plant Cell 19: 862–876. </ref>
<ref name="ref6">Bhalla N, Wynne DJ, Jantsch V, Dernburg AF (2008) ZHP-3 acts at crossovers to couple meiotic recombination with synaptonemal complex disassembly and bivalent formation in C. elegans. PLoS Genet 4: e1000235. </ref>
<ref name="ref7">Borner GV, Kleckner N, Hunter N (2004) Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell 117: 29–45. </ref>
<ref name="ref8"> Mercier R, Jolivet S, Vezon D, Huppe E, Chelysheva L, et al. (2005) Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not. Curr Biol 15: 692–701. </ref>
<ref name="ref9">Higgins JD, Armstrong SJ, Franklin FC, Jones GH (2004) The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis. Genes & Dev 18: 2557–2570. </ref>
<ref name="ref10">Wang K, Tang D, Wang M, Lu J, Yu H, et al. (2009) MER3 is required for normal meiotic crossover formation, but not for presynaptic alignment in rice. J Cell Sci 122: 2055–2063. </ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

Os02g0232100

2014-06-04T09:07:14Z

Simon: /* Mutation */

Please input one-sentence summary here.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.
===Mutation===
[[File:Figure111.jpg|right|thumb|250px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref3" />).'']]
60Co γ-ray irradiation was used to induce a sterile mutant from the japonica rice variety Wuxiangjing 9.The mutant showed normal vegetative growth but exhibited complete sterility (Figure 1A and 1B). Cytological observation of anthers showed that almost all pollens were shrunken and inviable (Figure 1C and 1D). When pollinated with wild-type (WT) pollens, the mutant spikelets were unable to set any seeds, suggesting that the female gametes were also sterile. The progenies of the heterozygous plants segregated from normal to sterile phenotype in a 3:1 ratio (fertile, 30; sterile, 10), indicating that a single recessive gene is responsible for the sterile phenotype<ref name="ref3" />.

===Expression（Mutant VS Wild type）===
[[File:Figure2.jpg|right|thumb|250px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure3.jpg|right|thumb|250px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure4.jpg|right|thumb|250px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.

===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.

==Labs working on this gene==
State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

==References==
<references>
<ref name="ref1"> Toby GG, Gherraby W, Coleman TR, Golemis EA (2003) A novel RING finger protein, human enhancer of invasion 10, alters mitotic progression through regulation of cyclin B levels. Mol Cell Biol 23: 2109–2122. </ref>
<ref name="ref2"> Singh MK, Nicolas E, Gherraby W, Dadke D, Lessin S, et al. (2007) HEI10 negatively regulates cell invasion by inhibiting cyclin B/Cdk1 and other promotility proteins. Oncogene 26: 4825–4832. </ref>
<ref name="ref3">Wang, K., Wang, M., Tang, D., Shen, Y., Miao, C., Hu, Q., ... & Cheng, Z. (2012).The role of rice HEI10 in the formation of meiotic crossovers. PLoS genetics, 8(7), e1002809. </ref>
<ref name="ref4">de Boer E, Stam P, Dietrich AJ, Pastink A, Heyting C (2006) Two levels of interference in mouse meiotic recombination. Proc Natl Acad Sci U S A 103: 9607–9612. </ref>
<ref name="ref5">Lhuissier FG, Offenberg HH, Wittich PE, Vischer NO, Heyting C (2007) The mismatch repair protein MLH1 marks a subset of strongly interfering crossovers in tomato. Plant Cell 19: 862–876. </ref>
<ref name="ref6">Bhalla N, Wynne DJ, Jantsch V, Dernburg AF (2008) ZHP-3 acts at crossovers to couple meiotic recombination with synaptonemal complex disassembly and bivalent formation in C. elegans. PLoS Genet 4: e1000235. </ref>
<ref name="ref7">Borner GV, Kleckner N, Hunter N (2004) Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell 117: 29–45. </ref>
<ref name="ref8"> Mercier R, Jolivet S, Vezon D, Huppe E, Chelysheva L, et al. (2005) Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not. Curr Biol 15: 692–701. </ref>
<ref name="ref9">Higgins JD, Armstrong SJ, Franklin FC, Jones GH (2004) The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis. Genes & Dev 18: 2557–2570. </ref>
<ref name="ref10">Wang K, Tang D, Wang M, Lu J, Yu H, et al. (2009) MER3 is required for normal meiotic crossover formation, but not for presynaptic alignment in rice. J Cell Sci 122: 2055–2063. </ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

Os02g0232100

2014-06-04T08:56:20Z

Simon: /* Knowledge Extension */

Please input one-sentence summary here.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.
===Mutation===
[[File:Figure1.jpg|right|thumb|250px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref3" />).'']]
60Co γ-ray irradiation was used to induce a sterile mutant from the japonica rice variety Wuxiangjing 9.The mutant showed normal vegetative growth but exhibited complete sterility (Figure 1A and 1B). Cytological observation of anthers showed that almost all pollens were shrunken and inviable (Figure 1C and 1D). When pollinated with wild-type (WT) pollens, the mutant spikelets were unable to set any seeds, suggesting that the female gametes were also sterile. The progenies of the heterozygous plants segregated from normal to sterile phenotype in a 3:1 ratio (fertile, 30; sterile, 10), indicating that a single recessive gene is responsible for the sterile phenotype<ref name="ref3" />.

===Expression（Mutant VS Wild type）===
[[File:Figure2.jpg|right|thumb|250px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure3.jpg|right|thumb|250px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure4.jpg|right|thumb|250px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.

===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.

==Labs working on this gene==
State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

==References==
<references>
<ref name="ref1"> Toby GG, Gherraby W, Coleman TR, Golemis EA (2003) A novel RING finger protein, human enhancer of invasion 10, alters mitotic progression through regulation of cyclin B levels. Mol Cell Biol 23: 2109–2122. </ref>
<ref name="ref2"> Singh MK, Nicolas E, Gherraby W, Dadke D, Lessin S, et al. (2007) HEI10 negatively regulates cell invasion by inhibiting cyclin B/Cdk1 and other promotility proteins. Oncogene 26: 4825–4832. </ref>
<ref name="ref3">Wang, K., Wang, M., Tang, D., Shen, Y., Miao, C., Hu, Q., ... & Cheng, Z. (2012).The role of rice HEI10 in the formation of meiotic crossovers. PLoS genetics, 8(7), e1002809. </ref>
<ref name="ref4">de Boer E, Stam P, Dietrich AJ, Pastink A, Heyting C (2006) Two levels of interference in mouse meiotic recombination. Proc Natl Acad Sci U S A 103: 9607–9612. </ref>
<ref name="ref5">Lhuissier FG, Offenberg HH, Wittich PE, Vischer NO, Heyting C (2007) The mismatch repair protein MLH1 marks a subset of strongly interfering crossovers in tomato. Plant Cell 19: 862–876. </ref>
<ref name="ref6">Bhalla N, Wynne DJ, Jantsch V, Dernburg AF (2008) ZHP-3 acts at crossovers to couple meiotic recombination with synaptonemal complex disassembly and bivalent formation in C. elegans. PLoS Genet 4: e1000235. </ref>
<ref name="ref7">Borner GV, Kleckner N, Hunter N (2004) Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell 117: 29–45. </ref>
<ref name="ref8"> Mercier R, Jolivet S, Vezon D, Huppe E, Chelysheva L, et al. (2005) Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not. Curr Biol 15: 692–701. </ref>
<ref name="ref9">Higgins JD, Armstrong SJ, Franklin FC, Jones GH (2004) The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis. Genes & Dev 18: 2557–2570. </ref>
<ref name="ref10">Wang K, Tang D, Wang M, Lu J, Yu H, et al. (2009) MER3 is required for normal meiotic crossover formation, but not for presynaptic alignment in rice. J Cell Sci 122: 2055–2063. </ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

Os02g0232100

2014-06-04T08:54:48Z

Simon: /* Evolution */

Please input one-sentence summary here.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.
===Mutation===
[[File:Figure1.jpg|right|thumb|250px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref3" />).'']]
60Co γ-ray irradiation was used to induce a sterile mutant from the japonica rice variety Wuxiangjing 9.The mutant showed normal vegetative growth but exhibited complete sterility (Figure 1A and 1B). Cytological observation of anthers showed that almost all pollens were shrunken and inviable (Figure 1C and 1D). When pollinated with wild-type (WT) pollens, the mutant spikelets were unable to set any seeds, suggesting that the female gametes were also sterile. The progenies of the heterozygous plants segregated from normal to sterile phenotype in a 3:1 ratio (fertile, 30; sterile, 10), indicating that a single recessive gene is responsible for the sterile phenotype<ref name="ref3" />.

===Expression（Mutant VS Wild type）===
[[File:Figure2.jpg|right|thumb|250px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure3.jpg|right|thumb|250px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure4.jpg|right|thumb|250px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.

===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.

===Knowledge Extension===

In hei10 mutants, only about 31% chiasmata were maintained. This number is similar to that of mer3 mutants, in which about 28% chiasmata were remained. Previous studies revealed that there are at least two classes of CO which occur in budding yeast and Arabidopsis <ref name="ref7"><ref name="ref8"><ref name="ref9">. Investigations of mer3 mutants in rice showed that apart from a reduced number of chiasmata, the remaining chiasmata distributed randomly among cells, suggesting that rice might also have at least two kinds of COs: one that appears to be sensitive to interference, whereas the other one is not <ref name="ref10">. The close number of residual chiasmata indicates similar contributions of HEI10 and MER3 during CO formation. Considering the close number and similar random distribution of residual chiasmata noted in mer3 and hei10 mutants, it is likely that HEI10 and MER3 may act in the same CO pathway, namely an interference-sensitive pathway.Besides a reduced chiasma frequency, hei10 also showed a precocious segregation of sister chromatids of some univalents at anaphase I.

==Labs working on this gene==
State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

==References==
<references>
<ref name="ref1"> Toby GG, Gherraby W, Coleman TR, Golemis EA (2003) A novel RING finger protein, human enhancer of invasion 10, alters mitotic progression through regulation of cyclin B levels. Mol Cell Biol 23: 2109–2122. </ref>
<ref name="ref2"> Singh MK, Nicolas E, Gherraby W, Dadke D, Lessin S, et al. (2007) HEI10 negatively regulates cell invasion by inhibiting cyclin B/Cdk1 and other promotility proteins. Oncogene 26: 4825–4832. </ref>
<ref name="ref3">Wang, K., Wang, M., Tang, D., Shen, Y., Miao, C., Hu, Q., ... & Cheng, Z. (2012).The role of rice HEI10 in the formation of meiotic crossovers. PLoS genetics, 8(7), e1002809. </ref>
<ref name="ref4">de Boer E, Stam P, Dietrich AJ, Pastink A, Heyting C (2006) Two levels of interference in mouse meiotic recombination. Proc Natl Acad Sci U S A 103: 9607–9612. </ref>
<ref name="ref5">Lhuissier FG, Offenberg HH, Wittich PE, Vischer NO, Heyting C (2007) The mismatch repair protein MLH1 marks a subset of strongly interfering crossovers in tomato. Plant Cell 19: 862–876. </ref>
<ref name="ref6">Bhalla N, Wynne DJ, Jantsch V, Dernburg AF (2008) ZHP-3 acts at crossovers to couple meiotic recombination with synaptonemal complex disassembly and bivalent formation in C. elegans. PLoS Genet 4: e1000235. </ref>
<ref name="ref7">Borner GV, Kleckner N, Hunter N (2004) Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell 117: 29–45. </ref>
<ref name="ref8"> Mercier R, Jolivet S, Vezon D, Huppe E, Chelysheva L, et al. (2005) Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not. Curr Biol 15: 692–701. </ref>
<ref name="ref9">Higgins JD, Armstrong SJ, Franklin FC, Jones GH (2004) The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis. Genes & Dev 18: 2557–2570. </ref>
<ref name="ref10">Wang K, Tang D, Wang M, Lu J, Yu H, et al. (2009) MER3 is required for normal meiotic crossover formation, but not for presynaptic alignment in rice. J Cell Sci 122: 2055–2063. </ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

Os02g0232100

2014-06-04T08:54:01Z

Simon: /* Function */

Please input one-sentence summary here.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.
===Mutation===
[[File:Figure1.jpg|right|thumb|250px|'' The rice hei10-1 mutant phenotype (from reference <ref name="ref3" />).'']]
60Co γ-ray irradiation was used to induce a sterile mutant from the japonica rice variety Wuxiangjing 9.The mutant showed normal vegetative growth but exhibited complete sterility (Figure 1A and 1B). Cytological observation of anthers showed that almost all pollens were shrunken and inviable (Figure 1C and 1D). When pollinated with wild-type (WT) pollens, the mutant spikelets were unable to set any seeds, suggesting that the female gametes were also sterile. The progenies of the heterozygous plants segregated from normal to sterile phenotype in a 3:1 ratio (fertile, 30; sterile, 10), indicating that a single recessive gene is responsible for the sterile phenotype<ref name="ref3" />.

===Expression（Mutant VS Wild type）===
[[File:Figure2.jpg|right|thumb|250px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure3.jpg|right|thumb|250px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure4.jpg|right|thumb|250px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.

===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.

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

==Labs working on this gene==
State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

==References==
<references>
<ref name="ref1"> Toby GG, Gherraby W, Coleman TR, Golemis EA (2003) A novel RING finger protein, human enhancer of invasion 10, alters mitotic progression through regulation of cyclin B levels. Mol Cell Biol 23: 2109–2122. </ref>
<ref name="ref2"> Singh MK, Nicolas E, Gherraby W, Dadke D, Lessin S, et al. (2007) HEI10 negatively regulates cell invasion by inhibiting cyclin B/Cdk1 and other promotility proteins. Oncogene 26: 4825–4832. </ref>
<ref name="ref3">Wang, K., Wang, M., Tang, D., Shen, Y., Miao, C., Hu, Q., ... & Cheng, Z. (2012).The role of rice HEI10 in the formation of meiotic crossovers. PLoS genetics, 8(7), e1002809. </ref>
<ref name="ref4">de Boer E, Stam P, Dietrich AJ, Pastink A, Heyting C (2006) Two levels of interference in mouse meiotic recombination. Proc Natl Acad Sci U S A 103: 9607–9612. </ref>
<ref name="ref5">Lhuissier FG, Offenberg HH, Wittich PE, Vischer NO, Heyting C (2007) The mismatch repair protein MLH1 marks a subset of strongly interfering crossovers in tomato. Plant Cell 19: 862–876. </ref>
<ref name="ref6">Bhalla N, Wynne DJ, Jantsch V, Dernburg AF (2008) ZHP-3 acts at crossovers to couple meiotic recombination with synaptonemal complex disassembly and bivalent formation in C. elegans. PLoS Genet 4: e1000235. </ref>
<ref name="ref7">Borner GV, Kleckner N, Hunter N (2004) Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell 117: 29–45. </ref>
<ref name="ref8"> Mercier R, Jolivet S, Vezon D, Huppe E, Chelysheva L, et al. (2005) Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not. Curr Biol 15: 692–701. </ref>
<ref name="ref9">Higgins JD, Armstrong SJ, Franklin FC, Jones GH (2004) The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis. Genes & Dev 18: 2557–2570. </ref>
<ref name="ref10">Wang K, Tang D, Wang M, Lu J, Yu H, et al. (2009) MER3 is required for normal meiotic crossover formation, but not for presynaptic alignment in rice. J Cell Sci 122: 2055–2063. </ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

Os02g0232100

2014-06-04T08:52:48Z

Simon: /* References */

Please input one-sentence summary here.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.

===Expression（Mutant VS Wild type）===
[[File:Figure2.jpg|right|thumb|250px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure3.jpg|right|thumb|250px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure4.jpg|right|thumb|250px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.

===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.

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

==Labs working on this gene==
State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

==References==
<references>
<ref name="ref1"> Toby GG, Gherraby W, Coleman TR, Golemis EA (2003) A novel RING finger protein, human enhancer of invasion 10, alters mitotic progression through regulation of cyclin B levels. Mol Cell Biol 23: 2109–2122. </ref>
<ref name="ref2"> Singh MK, Nicolas E, Gherraby W, Dadke D, Lessin S, et al. (2007) HEI10 negatively regulates cell invasion by inhibiting cyclin B/Cdk1 and other promotility proteins. Oncogene 26: 4825–4832. </ref>
<ref name="ref3">Wang, K., Wang, M., Tang, D., Shen, Y., Miao, C., Hu, Q., ... & Cheng, Z. (2012).The role of rice HEI10 in the formation of meiotic crossovers. PLoS genetics, 8(7), e1002809. </ref>
<ref name="ref4">de Boer E, Stam P, Dietrich AJ, Pastink A, Heyting C (2006) Two levels of interference in mouse meiotic recombination. Proc Natl Acad Sci U S A 103: 9607–9612. </ref>
<ref name="ref5">Lhuissier FG, Offenberg HH, Wittich PE, Vischer NO, Heyting C (2007) The mismatch repair protein MLH1 marks a subset of strongly interfering crossovers in tomato. Plant Cell 19: 862–876. </ref>
<ref name="ref6">Bhalla N, Wynne DJ, Jantsch V, Dernburg AF (2008) ZHP-3 acts at crossovers to couple meiotic recombination with synaptonemal complex disassembly and bivalent formation in C. elegans. PLoS Genet 4: e1000235. </ref>
<ref name="ref7">Borner GV, Kleckner N, Hunter N (2004) Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell 117: 29–45. </ref>
<ref name="ref8"> Mercier R, Jolivet S, Vezon D, Huppe E, Chelysheva L, et al. (2005) Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not. Curr Biol 15: 692–701. </ref>
<ref name="ref9">Higgins JD, Armstrong SJ, Franklin FC, Jones GH (2004) The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis. Genes & Dev 18: 2557–2570. </ref>
<ref name="ref10">Wang K, Tang D, Wang M, Lu J, Yu H, et al. (2009) MER3 is required for normal meiotic crossover formation, but not for presynaptic alignment in rice. J Cell Sci 122: 2055–2063. </ref>
</references>

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

Os02g0232100

2014-06-04T08:52:17Z

Simon: /* Labs working on this gene */

Please input one-sentence summary here.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.

===Expression（Mutant VS Wild type）===
[[File:Figure2.jpg|right|thumb|250px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure3.jpg|right|thumb|250px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure4.jpg|right|thumb|250px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.

===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.

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

==Labs working on this gene==
State Key Laboratory of Plant Genomics and Center for Plant Gene Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing,China.
Biotechnology Research Institute/National Key Facility for Gene Resources and Gene Improvement, Chinese Academy of Agricultural Sciences, Beijing, China.

==References==
Please input cited references here.

==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

Os02g0232100

2014-06-04T08:51:49Z

Simon: /* Evolution */

Please input one-sentence summary here.

==Annotated Information==
HEI10 exerts its effects in the recombination process through modification of diverse meiotic components during rice meiosis.
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.

===Expression（Mutant VS Wild type）===
[[File:Figure2.jpg|right|thumb|250px|'' Organization of the HEI10 gene and protein alignment (from reference <ref name="ref3" />).'']]
[[File:Figure3.jpg|right|thumb|250px|'' Meiosis in the hei10-1 mutant (from reference <ref name="ref3" />).'']]
[[File:Figure4.jpg|right|thumb|250px|'' Analysis of the distribution of HEI10 bright foci in WT
meiocytes (from reference <ref name="ref3" />).'']]

Using sterile plants that segregated in F2 and F3 populations, the gene was mapped on the long arm of rice chromosome 2, which was further narrowed to a 100-kb region. All genes within this region were amplified and sequenced. A single nucleotide G to A substitution was found at position 140 of the first exon of the Os02g0232100 gene (Figure 2). This substitution introduced a new translation initiation site (ATG) in the 5’-UTR, which would theoretically express a totally different peptide<ref name="ref3" />.
The hei10 mutant chromosomes behaved normally during leptotene and zygotene. Fully aligned chromosomes were detected during pachytene (Figure 3A). However, during diakinesis, the mutant cells showed a mixture of both univalent and bivalent chromosomes (Figure 3B). At metaphase I, the bivalents aligned well on the equatorial plate while some of the univalents were scattered in the nucleus (Figure 3C). In anaphase I, the bivalents separated normally but the scattered univalents segregated randomly. Besides those randomly distributed univalents, many univalents also aligned on the equatorial plate in metaphase I (Figure 3D) and underwent precocious separation of sister chromatids in anaphase I (Figure 3E), indicating a bipolar orientation of sister kinetochores. In telophase I and prophase II, an uneven number of chromosomes was observed in the two related cells. After the second division, tetrads with aberrant numbers of chromosomes were formed. In addition, multiple micronuclei were frequently observed (Figure 3F) <ref name="ref3" />.
To accurately define the spatial and temporal distribution of HEI10 during meiosis in rice, dual immunolocalization experiments were performed using polyclonal antibodies against REC8 and HEI10 protein, raised in rabbit and mouse, respectively. To obtain the precise localization of HEI10 bright foci on the chromosome, observation of scattered chromosomes at late pachytene revealed that one, two or three, frequently two, prominent foci localized on each pair of homologs (Figure 4A). Additionally, when two or three foci occurred on the same chromosome, they tended to be spaced far apart. Immunostaining using antibodies against HEI10 and CENH3 also revealed that about 95.6% HEI10 foci located outside CENH3 position at late pachytene (Figure 4B). To further explore whether those HEI10 bright foci were randomly distributed along bivalents, we measured the interfocus distance among bright HEI10 focus on the shortest chromosome of the cell and estimated the existence of interference using the interference parameter n of the gamma model<ref name="ref4" /><ref name="ref5" /> (Figure 4D). The result showed that HEI10 bright foci on a single chromosome displayed strong interference<ref name="ref3" />.

===Evolution===
HEI10 is the most likely ortholog of ZHP-3 and Zip3 in rice, although other proteins belonging to the Zip3 family cannot be excluded. In contrast to Zip3, both HEI10 and ZHP-3 exhibit a similar dynamic localization pattern, implying that the proteins may evolve new functions during meiotic recombination in multicellular organism. In C. elegans ZHP-3 was required for SC asymmetrical disassembly and normal bivalent structure <ref name="ref6" />. Such defects were not observed in rice, implying probable diversification of Zip3 homologs even among multicelluar organisms.

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 = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]

Os02g0232100

2014-06-04T08:49:52Z

Simon: /* Expression */

Os02g0232100

2014-06-04T08:47:20Z

Simon: /* Annotated Information */

Os02g0232100

2014-06-04T08:46:49Z

Simon: /* Function */

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==Annotated Information==
===Function===
Human Enhancer of Invasion 10 (HEI10; also known asCCNB1IP1) was first isolated from humans and it was demonstrated that the molecule played a role in the mitotic cell cycle <ref name="ref1" />. HEI10 functions as an E3 Ubiquitin ligase to regulate cell migration and invasion <ref name="ref1" /><ref name="ref2" />. Further studies in mice revealed that mutation of HEI10 led to dramatic meiosis defects, indicating an important role of HEI10 during meiosis. It was shown in mice that mutation of HEI10 resulted in high prevalence of univalent chromosomes during metaphase I, which finally leads to a sterile phenotype. Furthermore, results obtained in experiments using a yeast two-hybrid system suggest a function for HEI10 as E3 SUMO ligase in addition to the ubiquitin ligase role reported in somatic cells<ref name="ref2" />.It is further known that HEI10 is required for meiotic CO formation. Consistent with this, it suggests that the role of HEI10 in rice might be the homolog of budding yeast Zip3 and C. elegans ZHP-3. Those genes may play both conserved and divergent roles in homologous recombination in their respective species. It suggests that also in rice HEI10 is essential for reciprocal recombination between homologous chromosomes<ref name="ref3" />.

===Expression===
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===Evolution===
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==Labs working on this gene==
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==References==
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==Structured Information==
{{JaponicaGene|
GeneName = Os02g0232100|
Description = Zinc finger, RING-type domain containing protein|
Version = NM_001052906.1 GI:115445182 GeneID:4328810|
Length = 3452 bp|
Definition = Oryza sativa Japonica Group Os02g0232100, 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 2|Chromosome 2]]|
AP = Chromosome 2:7496475..7499926|
CDS = 7496552..7496823,7497026..7497189,7497285..7497540,7497756..7497786,7497862..7497907 ,7498216..7498288,7498408..7498477|
GCID = <gbrowseImage1>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage1>|
GSID = <gbrowseImage2>
name=NC_008395:7496475..7499926
source=RiceChromosome02
preset=GeneLocation
</gbrowseImage2>|
CDNA = <cdnaseq>atgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaagccatatgagacctgttgatacaaatccaaatgatgattggacaaatatgtcaatggctggagtttctccacagatacttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaa</cdnaseq>|
AA = <aaseq>MKCNACWRELEGQAVSTTCGHLLCTEDAKKILSNDAACPICDQV LSKSHMRPVDTNPNDDWTNMSMAGVSPQILMKSAYRSVMFYIGQKELEMQYKMNRIVG QCRQKCELMQAKFTEKLEEVHTAYQKMAKKCQLMEQEVENLSRDKQELQEKFAEKSRQ KRKLDEMYDQLRSEYESAKRSAIQPANNYFPRAQPDLFSGVPNIMDSSDPLRQGLAGL PETPGRRDEGWAPPPRQRRSTSGPFELSAGSPAHNAAPPVDIRPRQPARPVFGTAMNN TSAALRNMIISPVKRPQLSRNRPHMFT</aaseq>|
DNA = <dnaseqindica>3104..3375#2738..2901#2387..2642#2141..2171#2020..2065#1639..1711#1450..1519#gcaaagcaattttgccattctcccgtttcaaacccgcgctctctctctctctctctctctctccccctccctgccgctgctgccgccaccgccgcggcggcgccccgattcggtgatcccccgctcgcccatctccgccgtgacgtatcggcgacgcggagggagccccccaccccccacctccggatctagggtttcgaggcccaggtggcgttgcgctgcgcccccgctcctccgcctctcctccgccgccgagggttcgcgtgaggtaaccgacgcgcccccgtttggtaatcgcaccggcagatatcgtgggggttcgtttctccgtggcgtgttttggtgatggatccgatctaggtttgttttaggggctattttgacccaaatttcaggtgttttgcggagattgtagtgtttctacctagaaatactgcgattattggtgcgtttggagatgccactggctgtgaaatttcctgtgggaactcgtttccagttcgctggggatttatggaatggttacggttttgcaatattacgggttttgcgaactaaggcttcggtttggtcgctcatgggagcaatatttagtacgattgtcatcgccatttctgcaattccaccagcgattggttgctctgttcgtactcccaatagaaagcttgctccgctgctgatatttaaagtgcttcgttggtggatttcaactgcagatgttgggctctgaaggaaatggtgcaatttcttcgtgcatgatgctcgtatattgtgtctattatgtagtcctgcctgcttcggtggttctagatgctacagctttatctgccattctatttatgctgcattctatgttcagggcatgcgcaaaatattattgaaatgatcatttacttgaagataattggatttatttcattaattgctccgtaagccttgattcataggtagtattatgctttttttactactaagtctgtagaatcccatctctcctgcaatagctcataagaaagtgacatgttgtgcgatgcagaactggttctgaatgcatagtccttagtgcattttgagctaccctttgtcaggactgtagtctctgtaaatagtgggagaattcattggctaaaaagatgacaaaagctagctgcttatcagtattaagtgtcgcaccatttcttttatcatgcgagtagtgcaagtgcctgaagggttatcccatttagctattccaaatggcacatacgaaatatgattatttgttcatttgatccgtgatacttaggtagctatgatctgtctctgctcggtttgtttttctgttcagacatgcagaatacaactgctgtctgtattcagtttatatcttgcaactcataattctatgtttctacctgcaggtttctgcgtattaccagctctgtacttaatttactagtgagaagcaaaatgaagtgcaatgcttgctggcgggagttggaagggcaagctgtatcaacaacctgcggtcatcttttatgtatgcctgcaatcttccttactagttttgcttaaccagtttctcattgtaaactaacattgtctgctgctattgcagtatacataactgatccttgctgatgattatccgttgagcaggtacagaggatgctaagaaaatactgagcaatgatgctgcatgcccgatttgtgatcaagtgctttcaaaaaggtctgttgtttactcttgaagttgtattcattatctcatactacattttctcttctttcctgtaagctatttttgaatgtacatcatgatttgacaattagggatgccaaaagtacctcaaacgcatacatatttatcatctcgatggttattgccatgtcatttcatccgaatgggtgcttaacacatggagtttactttctagcatgctttgaagcttttatacttctttgatgtgttcgagtgaaaatggactccagtggtttactattttgagacagtctaaacattataattttttttctcagccatatgagacctgttgatacaaatccaaatgatgattggacaaatgtaagttcgcttacgattcattgtagataaaagcaatccttctttttctaacagccagtctgctcattggtacagatgtcaatggctggagtttctccacagatacgtatccttattattttctcattgtccgtaagattcttgtgctacttttgttgaaaatcatcactaactgctatatggaatctatggcgtagctagcctcttgtcctgttttattgtcatagtaatgcaaaaatctgaaagatcaaaattgtttatatgattattgtctaaggcttggtgactattcagattttccctaactatgatgctaatcagttatgaagagtgcatacagaagtgtcatgttttacattgggcaaaaggaactggagatgcagtacaagatgaacagaattgttggtcagtgtaggcaaaagtgtgaacttatgcaggcaaagttcactgagaagctggaagaagttcatactgcataccagaaaatggccaaaaaatgccagttgatggaacaagaggttgaaaacttgtcgagggataagcaagagctacaagaaaaatttgctgagaaatccaggtcagtatgtacagtacatttctagtttaattctcatatgaataaacctctgcttcatccaacttatatggagaattgactgctgtcatttccaggcagaaaaggaagcttgatgagatgtatgatcagctgagaagcgagtatgagtcggcaaagcgttcagcaattcaacctgcgaacaactacttcccaagagcccagccagacctgttctcaggcgtgcccaacataatggatagcagcgaccctctgagacaaggtaatttaacatgttatcattcagcttattagttctctcataccatataactcatcctttcctctgtgcacacccttttgttatgcttccaaccagaaaaactgttaactgtaatgcttggattgctagtagccactattaaatatataaaagttaaaatgcttttagccctccacaaaaacagcttattcactggctgcaggattggctggtcttcctgaaactccagggcggagagatgagggatgggctccaccaccaaggcaacgtcggtccacctctggaccatttgagctgtctgcgggatctcctgctcacaatgcggcgcctccggttgatattaggcccagacagccagcacggcccgtattcggcactgccatgaataatacttctgcagctctgcgaaatatgataatctcgcctgtgaaacgtcctcagctttcccgaaaccgtccacatatgttcacgtaatgtcctccttattgctcatgcatgtatttcacatctaaatgaagcagtttgggaatgaaaattgatgtccaacttgc</dnaseqindica>|
Link = [http://www.ncbi.nlm.nih.gov/nuccore/NM_001052906.1 RefSeq:Os02g0232100]|
}}
[[Category:Genes]]
[[Category:Japonica mRNA]]
[[Category:Oryza Sativa Japonica Group]]
[[Category:Japonica Genes]]
[[Category:Japonica Chromosome 2]]
[[Category:Chromosome 2]]