title: Comprehensive analyses of the annexin gene family in wheat.
abstract: Annexins are an evolutionarily conserved multigene family of calcium-dependent phospholipid binding proteins that play important roles in stress resistance and plant development. They have been relatively well characterized in model plants Arabidopsis (Arabidopsis thaliana) and rice (Oryza sativa), but nothing has been reported in hexaploid bread wheat (Triticum aestivum) and barely (Hordeum vulgare), which are the two most economically important plants.
  
  
  
  Based on available genomic and transcriptomic data, 25 and 11 putative annexin genes were found through in silico analysis in wheat and barley, respectively. Additionally, eight and 11 annexin genes were identified from the draft genome sequences of Triticum urartu and Aegilops tauschii, progenitor for the A and D genome of wheat, respectively. By phylogenetic analysis, annexins in these four species together with other monocots and eudicots were classified into six different orthologous groups. Pi values of each of Ann1-12 genes among T. aestivum, T. urartu, A. tauschii and H. vulgare species was very low, with the exception of Ann2 and Ann5 genes. Ann2 gene has been under positive selection, but Ann6 and Ann7 have been under purifying selection among the four species in their evolutionary histories. The nucleotide diversities of Ann1-12 genes in the four species were 0.52065, 0.59239, 0.60691 and 0.53421, respectively. No selective pressure was operated on annexin genes in the same species. Gene expression patterns obtained by real-time PCR and re-analyzing the public microarray data revealed differential temporal and spatial regulation of annexin genes in wheat under different abiotic stress conditions such as salinity, drought, cold and abscisic acid. Among those genes, TaAnn10 is specifically expressed in the anther but fails to be induced by low temperature in thermosensitive genic male sterile lines, suggesting that specific down-regulation of TaAnn10 is associated with conditional male sterility in wheat.
  
  
  
  This study analyzed the size and composition of the annexin gene family in wheat and barley, and investigated differential tissue-specific and stress responsive expression profiles of the gene family in wheat. These results provided significant information for understanding the diverse roles of plant annexins and opened a new avenue for functional studies of cold induced male sterility in wheat.
authors.affiliation: Beijing Engineering Research Center for Hybrid Wheat, The Municipal Key Laboratory of the Molecular Genetics of Hybrid Wheat, Beijing Academy of Agriculture and Forestry Sciences, Beijing, 100097, China.|College of Life Science, Capital Normal University, Beijing, 100048, China.
authors.forename: Lei
authors.initials: L
authors.lastname: Xu
authors.affiliation: Beijing Engineering Research Center for Hybrid Wheat, The Municipal Key Laboratory of the Molecular Genetics of Hybrid Wheat, Beijing Academy of Agriculture and Forestry Sciences, Beijing, 100097, China. tangyimiao@126.com.
authors.forename: Yimiao
authors.initials: Y
authors.lastname: Tang
authors.affiliation: Beijing Engineering Research Center for Hybrid Wheat, The Municipal Key Laboratory of the Molecular Genetics of Hybrid Wheat, Beijing Academy of Agriculture and Forestry Sciences, Beijing, 100097, China.
authors.forename: Shiqing
authors.initials: S
authors.lastname: Gao
authors.affiliation: College of Life Science, Hebei Normal University of Science and Technology, Qinhuangdao, 066600, China.
authors.forename: Shichao
authors.initials: S
authors.lastname: Su
authors.affiliation: Beijing Engineering Research Center for Hybrid Wheat, The Municipal Key Laboratory of the Molecular Genetics of Hybrid Wheat, Beijing Academy of Agriculture and Forestry Sciences, Beijing, 100097, China.
authors.forename: Lin
authors.initials: L
authors.lastname: Hong
authors.affiliation: Beijing Engineering Research Center for Hybrid Wheat, The Municipal Key Laboratory of the Molecular Genetics of Hybrid Wheat, Beijing Academy of Agriculture and Forestry Sciences, Beijing, 100097, China.
authors.forename: Weiwei
authors.initials: W
authors.lastname: Wang
authors.affiliation: Beijing Engineering Research Center for Hybrid Wheat, The Municipal Key Laboratory of the Molecular Genetics of Hybrid Wheat, Beijing Academy of Agriculture and Forestry Sciences, Beijing, 100097, China.
authors.forename: Zhaofeng
authors.initials: Z
authors.lastname: Fang
authors.affiliation: Beijing Engineering Research Center for Hybrid Wheat, The Municipal Key Laboratory of the Molecular Genetics of Hybrid Wheat, Beijing Academy of Agriculture and Forestry Sciences, Beijing, 100097, China.
authors.forename: Xueyin
authors.initials: X
authors.lastname: Li
authors.affiliation: Beijing Engineering Research Center for Hybrid Wheat, The Municipal Key Laboratory of the Molecular Genetics of Hybrid Wheat, Beijing Academy of Agriculture and Forestry Sciences, Beijing, 100097, China.
authors.forename: Jinxiu
authors.initials: J
authors.lastname: Ma
authors.affiliation: Beijing Engineering Research Center for Hybrid Wheat, The Municipal Key Laboratory of the Molecular Genetics of Hybrid Wheat, Beijing Academy of Agriculture and Forestry Sciences, Beijing, 100097, China.
authors.forename: Wei
authors.initials: W
authors.lastname: Quan
authors.affiliation: Beijing Engineering Research Center for Hybrid Wheat, The Municipal Key Laboratory of the Molecular Genetics of Hybrid Wheat, Beijing Academy of Agriculture and Forestry Sciences, Beijing, 100097, China.
authors.forename: Hui
authors.initials: H
authors.lastname: Sun
authors.affiliation: State Key Laboratory of Protein and Plant Gene Research, Peking-Tsinghua Center for Life Sciences, School of Advanced Agricultural Sciences and School of Life Sciences, Peking University, Beijing, 100871, China.
authors.forename: Xia
authors.initials: X
authors.lastname: Li
authors.affiliation: Beijing Engineering Research Center for Hybrid Wheat, The Municipal Key Laboratory of the Molecular Genetics of Hybrid Wheat, Beijing Academy of Agriculture and Forestry Sciences, Beijing, 100097, China.
authors.forename: Yongbo
authors.initials: Y
authors.lastname: Wang
authors.affiliation: Beijing Engineering Research Center for Hybrid Wheat, The Municipal Key Laboratory of the Molecular Genetics of Hybrid Wheat, Beijing Academy of Agriculture and Forestry Sciences, Beijing, 100097, China.
authors.forename: Xiangzheng
authors.initials: X
authors.lastname: Liao
authors.affiliation: Beijing Engineering Research Center for Hybrid Wheat, The Municipal Key Laboratory of the Molecular Genetics of Hybrid Wheat, Beijing Academy of Agriculture and Forestry Sciences, Beijing, 100097, China.
authors.forename: Jiangang
authors.initials: J
authors.lastname: Gao
authors.affiliation: Beijing Engineering Research Center for Hybrid Wheat, The Municipal Key Laboratory of the Molecular Genetics of Hybrid Wheat, Beijing Academy of Agriculture and Forestry Sciences, Beijing, 100097, China.
authors.forename: Fengting
authors.initials: F
authors.lastname: Zhang
authors.affiliation: State Key Laboratory of Protein and Plant Gene Research, Peking-Tsinghua Center for Life Sciences, School of Advanced Agricultural Sciences and School of Life Sciences, Peking University, Beijing, 100871, China. lei.li@pku.edu.cn.
authors.forename: Lei
authors.initials: L
authors.lastname: Li
authors.affiliation: Beijing Engineering Research Center for Hybrid Wheat, The Municipal Key Laboratory of the Molecular Genetics of Hybrid Wheat, Beijing Academy of Agriculture and Forestry Sciences, Beijing, 100097, China. cp_zhao@vip.sohu.com.
authors.forename: Changping
authors.initials: C
authors.lastname: Zhao
mesh_terms: D017302:Annexins / Q000235:genetics*; D019295:Computational Biology / Q000379:methods; D019143:Evolution, Molecular; D018506:Gene Expression Regulation, Plant; D017343:Genes, Plant; D018745:Genome, Plant; D023281:Genomics / Q000379:methods; D059014:High-Throughput Nucleotide Sequencing; D005810:Multigene Family; D009928:Organ Specificity / Q000235:genetics; D010802:Phylogeny; D013312:Stress, Physiological / Q000235:genetics; D014908:Triticum / Q000145:classification / Q000235:genetics*
chemical_list: D017302:Annexins
keywords: Annexin; Calcium signaling; Cold induction; Hybrid wheat; Phylogenetic analysis; Thermosensitive genic male sterile (TGMS)