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Annotated Information
Function
ETERNAL TAPETUM 1(EAT1), a basic helix-loop-helix transcription factor conserved in land
plants, positively regulates programmed cell death in tapetal cells in rice anthers. eat1 exhibits
delayed tapetal cell death and aborted pollen formation, causing complete male sterility[1].
EAT1 directly regulates the expression of OsAP25 and OsAP37, which encode aspartic proteases that induce programmed cell death. In addition, EAT1 can interact with the TAPETUM DEGENERATION RETARDATION (TDR)protein and acts downstream of TDR[1]. TDR is also a key factor in regulates programmed cell death in tapetal cells[2].
Mutation
There are three kinds of EAT1 Mutation show in figure 2. The mutant exhibits normal vegetative development and female organ formation, but is completely male sterile and has shrunken anthers and aborted pollen grains (figure 3).
The eat1-1 mutant anthers appeared to undergo normal meiosis, forming tetrads of haploid
microspores at late stage 8. But after stage 10, the eat1-1 mutant had thicker tapetal cells, and abnormal abortion of the anther locule microspores. The result of terminal deoxynucleotidyl transferase-mediated dUTP nick-end labeling (TUNEL) assay and transmission electron microscopy (TEM) suggesting eat1-1 mutant cause delayed and abnormal PCD in tapetal cell.
Expression
The result of qRT–PCR indicated that in the wild type, EAT1 is weakly expressed in roots, shoots and leaves, and highly expressed in the anther from stage 7 to 12, while a dramatic reduction in expression was detected in the anthers in all three eat1 alleles (Fig. 4a). In GUS stained EAT1pro:GUS transgenic flowers, GUS signals started to appear in anthers at stage 7, became stronger from stage 8 to 9, and were nearly undetectable at stage 12 (Fig. 4b). Further in situ RNA hybridization indicated that EAT1 is highly expressed in the tapetum (Fig. 4c).
Evolution
Using the full-length EAT1 protein sequence to search available public databases and retrieved a total of 26 homologues from 10 diverse plant species from moss, pteridophytes, to angiosperms. The result show that EAT1 and three homologues from Sorghum bicolour (Sb04g030850), Zea mays (ZmLOC100282922) and Brachypodium distachyon (BradXP_003580474), respectively, were grouped in a subclade. EAT1 has one homologue from rice (OsbHLH142), which shares 40.8% identity with EAT1 in the HLH and DUF domains, and three homologues from Arabidopsis (AtbHLH091, AtbHLH089, AtbHLH010), which share an average of B40% identity with EAT1 in these two conserved domains.
Knowledge Extension
Tapetal cells is the innermost anther wall layer. After the meiosis of microspore mother cells it enter into programmed cell death (PCD). This degradation process is essential for microspore development and provide materials for pollen wall maturation; as a result, premature or delayed tapetal degradation causes male sterility.
In rice Several transcriptional regulators have been reported to be associated with tapetal degeneration. GAMYB[3], UNDEVELOPED TAPETUM1[4], TAPETUM DEGENERATION RETARDATION(TDR)[2], PERSISTENT TAPETAL CELL 1 (PTC1)[5], and APOPTOSIS INHIBITOR5 (API5)[6], ETERNAL TAPETUM 1(EAT1)[1]. TDR, the rice ortholog of the Arabidopsis AMS protein[2]. PTC,the rice ortholog of the Arabidopsis MS1 protein[5]. EAT1 shares sequence similarity with AtbHLH089 and AtbHLH091[1,7].
GA is a kind of important plant hormone, plays an important role in the process of plant growth and development, in some experiment show that GA signal is necessary for entry of the tapetum into PCD[3,8].
Labs working on this gene
- State Key Laboratory of Hybrid Rice, School of Life Sciences and Biotechnology, Shanghai Jiao Tong University, Shanghai, China
- Division of Plant Sciences,School of Biosciences, University of Nottingham, Loughborough, Leics, UK
- Department of Energy Plant Research Laboratory, Michigan State University, East Lansing, Michigan, USA.
References
[1] Ningning Niu. et al. EAT1 promotes tapetal cell death by regulating aspartic proteases during male reproductive development in rice. NATURE COMMUNICATIONS. 4:1445. (2013). DOI: 10.1038/ncomms2396
[2] Hui Li. et al. PERSISTENT TAPETAL CELL1 encodes a PHD-finger protein that is required for tapetal cell death and pollen development in rice. Plant Physiol.
156, 615–630 (2011).
[3] Aya, K. et al. Gibberellin modulates anther development in rice via the transcriptional regulation of GAMYB. Plant cell 21 1453–1472 (2009).
[4] Jung, K. H. et al. Rice Undeveloped Tapetum1 is a major regulator of early tapetum development. Plant Cell 10, 2705–2222 (2005).
[5] Li, H. et al. PERSISTENT TAPETAL CELL1 encodes a PHD-finger protein that is required for tapetal cell death and pollen development in rice. Plant Physiol.
156, 615–630 (2011).
[6] Li, X. W. et al. Rice APOPTOSIS INHIBITOR5 coupled with two DEAD-box adenosine 5’-triphosphate-dependent RNA helicases regulates tapetum degeneration. Plant Cell 23, 1416–1434 (2011).
[7] Li, X. X. et al. Genome-wide analysis of basic/helix-loop-helix transcription factor family in rice and Arabidopsis. Plant Physiol. 141, 1167–1184 (2006).
[8] Andrew R.G. Plackett. et al. Gibberellin control of stamen development: a fertile field. cell. 16: 568-578 (2011).
Structured Information
| Gene Name
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Os04g0599300
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| Description
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Basic helix-loop-helix dimerisation region bHLH domain containing protein
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| Version
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NM_001060284.1 GI:115460297 GeneID:4336865
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| Length
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3433 bp
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| Definition
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Oryza sativa Japonica Group Os04g0599300, complete gene.
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| Source
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Oryza sativa Japonica Group
ORGANISM Oryza sativa Japonica Group
Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
clade; Ehrhartoideae; Oryzeae; Oryza.
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| Chromosome
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Chromosome 4
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| Location
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Chromosome 4:30633942..30637374; between XY409-4 and XY409-5
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| Sequence Coding Region
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30634155..30634250,30634354..30634773,30635587..30636465
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| Expression
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GEO Profiles:Os04g0599300
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| Genome Context
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<gbrowseImage1>
name=NC_008397:30633942..30637374
source=RiceChromosome04
preset=GeneLocation
</gbrowseImage1>
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| Gene Structure
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<gbrowseImage2>
name=NC_008397:30633942..30637374
source=RiceChromosome04
preset=GeneLocation
</gbrowseImage2>
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| Coding Sequence
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<cdnaseq>atgattgttggggctggttactttgaggattcccacgatcaaagtctcatggcaggatctttgatccatgactcaaatcaagctcctgcaagcagtgaaaacacaagcattgatttgcagaaattcaaagtgcacccgtactcaacagaagctctctcgaatacggccaatctagctgaagctgcaagagcaattaaccaccttcaacatcaactagaaattgatttggagcaagaggttcccccagtagaaactgcaaactgggatccagctatctgcactataccagatcatatcatcaaccatcagtttagcgaagatccacaaaacatattggtggagcaacagatccagcagtatgattctgcactttatccaaatggtgtttacacacctgcaccagatctccttaatcttatgcagtgcacaatggctccagcattcccggcaacgacatccgtattcggtgacacaacactgaatggtactaactatttggatcttaacggtgaacttacaggagtagcagcggttccagacagtgggagtgggttgatgtttgctagtgattcagctctccagttagggtaccatggtactcaatctcatctaataaaggatatctgccactcgttgccccaaaattatgggttgtttcccagtgaggacgaacgagatgtgattattggtgttggaagtggagatctttttcaggagatagatgacaggcagtttgatagtgtacttgaatgcaggagagggaagggtgagttcggaaagggcaagggaaaagctaattttgcaactgagagagagaggcgggagcagctaaatgtgaagttcaggaccctaagaatgctcttcccaaatcctaccaagaatgacagggcctcaatagtaggtgatgccattgagtatatagatgagctcaatcgaacagtgaaggagctgaagatcctggtggaacagaagaggcatggaaataacaggagaaaggtgttaaagttggatcaagaggcagccgctgatggcgagagctcatcgatgaggccagtgagggatgatcaagacaatcagctccatggagccataaggagctcatgggttcagaggaggtcaaaggaatgccacgttgatgtccgcatagtggacgatgaagtaaacatcaagctcactgaaaagaagaaggccaactctctgcttcatgcagcaaaggttctagatgagttccagctcgagcttatccatgtagtgggtgggattataggtgatcaccatatattcatgttcaacactaaggtatcagaaggttcggcggtttatgcatgtgcagtggcaaagaagctccttcaagcagtggacgtgcaacaccaggccctcgacatattcaactaa</cdnaseq>
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| Protein Sequence
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<aaseq>MIVGAGYFEDSHDQSLMAGSLIHDSNQAPASSENTSIDLQKFKV HPYSTEALSNTANLAEAARAINHLQHQLEIDLEQEVPPVETANWDPAICTIPDHIINH QFSEDPQNILVEQQIQQYDSALYPNGVYTPAPDLLNLMQCTMAPAFPATTSVFGDTTL NGTNYLDLNGELTGVAAVPDSGSGLMFASDSALQLGYHGTQSHLIKDICHSLPQNYGL FPSEDERDVIIGVGSGDLFQEIDDRQFDSVLECRRGKGEFGKGKGKANFATERERREQ LNVKFRTLRMLFPNPTKNDRASIVGDAIEYIDELNRTVKELKILVEQKRHGNNRRKVL KLDQEAAADGESSSMRPVRDDQDNQLHGAIRSSWVQRRSKECHVDVRIVDDEVNIKLT EKKKANSLLHAAKVLDEFQLELIHVVGGIIGDHHIFMFNTKVSEGSAVYACAVAKKLL QAVDVQHQALDIFN</aaseq>
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| Gene Sequence
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<dnaseqindica>3125..3220#2602..3021#910..1788#tttgacctttattatatggtcataaagacccttcagcaaaatgattgttactgctatcggcattttctgttgtttttcttttggaatcaatcttgtgtgacaccattgtattgtttcatgtcttgccactataatagtcttggcatagcactggatctcatgagtctttgagccgcaaattcatgaacataagttctttccattcaaccgttggctgaggcaaagatacaggtatgttttttccagtgcttgctactactgtttgcaggatgcaaatcctaattagcattggtttatgtttctgtaaattagttgttaagttctatagaactttcaatcatactgaatttacagttcttacttttagtgatcagcttataataaatgaagtatatttggcattggcaatgatttcaagctactcagcattttactgattaattagtaaacttggggtggttgaagcacattttatcaaacatcaatatgaatatgattagaggcaaagaaagatggtaaggagtttgttaggtctgcaacaagcaaagttgcttcatgtctcattaatcatgctatatgcaacttctctacacggaataaacagacagacagattgcgtagcttaaactccacggctccatcttcccttgaaacaaccaaaacagctaagccaactgaaaattttcatgtccgattgaattatatccactgcttcattcatgttgagtagccctgtttcccttaatatgtgcattgcaagtaatttctattttagcactagattagcacccatctaagatgctatttgtccttcattttcatcctgtccttgattcttctgctcatatgtttttttacttgtgttggttttagatttggagcgaaggtgcctagcactgttttgccaaaatgattgttggggctggttactttgaggattcccacgatcaaagtctcatggcaggatctttgatccatgactcaaatcaagctcctgcaagcagtgaaaacacaagcattgatttgcagaaattcaaagtgcacccgtactcaacagaagctctctcgaatacggccaatctagctgaagctgcaagagcaattaaccaccttcaacatcaactagaaattgatttggagcaagaggttcccccagtagaaactgcaaactgggatccagctatctgcactataccagatcatatcatcaaccatcagtttagcgaagatccacaaaacatattggtggagcaacagatccagcagtatgattctgcactttatccaaatggtgtttacacacctgcaccagatctccttaatcttatgcagtgcacaatggctccagcattcccggcaacgacatccgtattcggtgacacaacactgaatggtactaactatttggatcttaacggtgaacttacaggagtagcagcggttccagacagtgggagtgggttgatgtttgctagtgattcagctctccagttagggtaccatggtactcaatctcatctaataaaggatatctgccactcgttgccccaaaattatgggttgtttcccagtgaggacgaacgagatgtgattattggtgttggaagtggagatctttttcaggagatagatgacaggcagtttgatagtgtacttgaatgcaggagagggaagggtgagttcggaaagggcaagggaaaagctaattttgcaactgagagagagaggcgggagcagctaaatgtgaagttcaggaccctaagaatgctcttcccaaatcctaccaaggttagtcttattcatcatcttgcaagttattagttgtttaggctgtaaataacttggtgattctcacattaacagacaaccactcagattttcaataatatttccatttgttactcatgctctgaagataatcaaaattttaaatatcctcatccatttattctcagagaactaatgattcaaaaactgccaacaccaatatagctccggtttagcaaatctctgtttttttacagatcacaaatacctaacagtaaatttataagtctgtgtattcatctaactggtataaattttgaaattatctgtccaaaatttcttcaagttgcgttaccacattttgatgcatatgtatatggaatatgctgtctgatatatcactcaacatgattgttttttgaaaaatagttcatcagtatgatgttctttactgataacagtgccatgttattaagggttgttttggttttaagccaaattatgccctaccaaattgttggcattttgaaaagttatttggcaaagtttggcttgccaccaaagttggtcaagttttggcactaccaatatattgacatggtaacaaatcaaaacacccctaattgtgttcatcctaaccaagtgagttagcccttctagttagctaggagaaagcaatagaagcattcagttcgatatttcctatgttccttgccttttttgtgtgttagcacattccacaatgttatcatcctcatgatgttacccttcaacaagattgtagcacttaaatatcttggttgtggcactaatgtgttacaaactgtgctctagaatgacagggcctcaatagtaggtgatgccattgagtatatagatgagctcaatcgaacagtgaaggagctgaagatcctggtggaacagaagaggcatggaaataacaggagaaaggtgttaaagttggatcaagaggcagccgctgatggcgagagctcatcgatgaggccagtgagggatgatcaagacaatcagctccatggagccataaggagctcatgggttcagaggaggtcaaaggaatgccacgttgatgtccgcatagtggacgatgaagtaaacatcaagctcactgaaaagaagaaggccaactctctgcttcatgcagcaaaggttctagatgagttccagctcgagcttatccatgtagtgggtgggattataggtgatcaccatatattcatgttcaacactaaggtaagtaacaattcagttttcttaaagtagaatcaaagattctttttgtcccattacacatgttagcatcgatagtaacgattcatcatccatggcaactcaggtatcagaaggttcggcggtttatgcatgtgcagtggcaaagaagctccttcaagcagtggacgtgcaacaccaggccctcgacatattcaactaatctttagcaacagtactgattatctgaacaatgtcctagattttcagttaccttgctgagcaaacttatttgaccaggattggagagaattttatctttagcactagctacctagcaaaacttcttaacaatttggccatgtaacggcttgctgctgtccggttgtacaccttaactagcctgactaggaaagctttgatgcttgtcttgtgt</dnaseqindica>
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| External Link(s)
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NCBI Gene:Os04g0599300, RefSeq:Os04g0599300
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