title: IRREGULAR POLLEN EXINE1 Is a Novel Factor in Anther Cuticle and Pollen Exine Formation.
abstract: Anther cuticle and pollen exine are protective barriers for pollen development and fertilization. Despite that several regulators have been identified for anther cuticle and pollen exine development in rice (Oryza sativa) and Arabidopsis (Arabidopsis thaliana), few genes have been characterized in maize (Zea mays) and the underlying regulatory mechanism remains elusive. Here, we report a novel male-sterile mutant in maize, irregular pollen exine1 (ipe1), which exhibited a glossy outer anther surface, abnormal Ubisch bodies, and defective pollen exine. Using map-based cloning, the IPE1 gene was isolated as a putative glucose-methanol-choline oxidoreductase targeted to the endoplasmic reticulum. Transcripts of IPE1 were preferentially accumulated in the tapetum during the tetrad and early uninucleate microspore stage. A biochemical assay indicated that ipe1 anthers had altered constituents of wax and a significant reduction of cutin monomers and fatty acids. RNA sequencing data revealed that genes implicated in wax and flavonoid metabolism, fatty acid synthesis, and elongation were differentially expressed in ipe1 mutant anthers. In addition, the analysis of transfer DNA insertional lines of the orthologous gene in Arabidopsis suggested that IPE1 and their orthologs have a partially conserved function in male organ development. Our results showed that IPE1 participates in the putative oxidative pathway of C16/C18 ω-hydroxy fatty acids and controls anther cuticle and pollen exine development together with MALE STERILITY26 and MALE STERILITY45 in maize.
authors.affiliation: National Maize Improvement Center of China, Beijing Key Laboratory of Crop Genetic Improvement, MOE Key Laboratory of Crop Heterosis and Utilization (X.C., H.S., Z.D., W.J.), and Department of Vegetable Sciences, Beijing Key Laboratory of Growth and Developmental Regulation for Protected Vegetable Crops (S.Y.), China Agricultural University, Beijing 100193, China.|State Key Laboratory of Plant Cell and Chromosome Engineering, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China (H.Z., L.Z., C.X., S.L., H.C.).|College of Agronomy, Southern Regional Collaborative Innovation Center for Grain and Oil Crops, Hunan Agricultural University, Changsha, Hunan 410128, China (H.L.).|Shanghai Center for Plant Stress Biology, Shanghai Institute for Biological Sciences, Chinese Academy of Sciences, Shanghai 201602, China (C.Z., R.L.); and.|University of the Chinese Academy of Sciences, Beijing 100039, China (H.Z.).
authors.forename: Xiaoyang
authors.initials: X
authors.lastname: Chen
authors.affiliation: National Maize Improvement Center of China, Beijing Key Laboratory of Crop Genetic Improvement, MOE Key Laboratory of Crop Heterosis and Utilization (X.C., H.S., Z.D., W.J.), and Department of Vegetable Sciences, Beijing Key Laboratory of Growth and Developmental Regulation for Protected Vegetable Crops (S.Y.), China Agricultural University, Beijing 100193, China.|State Key Laboratory of Plant Cell and Chromosome Engineering, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China (H.Z., L.Z., C.X., S.L., H.C.).|College of Agronomy, Southern Regional Collaborative Innovation Center for Grain and Oil Crops, Hunan Agricultural University, Changsha, Hunan 410128, China (H.L.).|Shanghai Center for Plant Stress Biology, Shanghai Institute for Biological Sciences, Chinese Academy of Sciences, Shanghai 201602, China (C.Z., R.L.); and.|University of the Chinese Academy of Sciences, Beijing 100039, China (H.Z.).
authors.forename: Hua
authors.initials: H
authors.lastname: Zhang
authors.affiliation: National Maize Improvement Center of China, Beijing Key Laboratory of Crop Genetic Improvement, MOE Key Laboratory of Crop Heterosis and Utilization (X.C., H.S., Z.D., W.J.), and Department of Vegetable Sciences, Beijing Key Laboratory of Growth and Developmental Regulation for Protected Vegetable Crops (S.Y.), China Agricultural University, Beijing 100193, China.|State Key Laboratory of Plant Cell and Chromosome Engineering, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China (H.Z., L.Z., C.X., S.L., H.C.).|College of Agronomy, Southern Regional Collaborative Innovation Center for Grain and Oil Crops, Hunan Agricultural University, Changsha, Hunan 410128, China (H.L.).|Shanghai Center for Plant Stress Biology, Shanghai Institute for Biological Sciences, Chinese Academy of Sciences, Shanghai 201602, China (C.Z., R.L.); and.|University of the Chinese Academy of Sciences, Beijing 100039, China (H.Z.).
authors.forename: Huayue
authors.identifier: 0000-0002-4656-3189
authors.initials: H
authors.lastname: Sun
authors.affiliation: National Maize Improvement Center of China, Beijing Key Laboratory of Crop Genetic Improvement, MOE Key Laboratory of Crop Heterosis and Utilization (X.C., H.S., Z.D., W.J.), and Department of Vegetable Sciences, Beijing Key Laboratory of Growth and Developmental Regulation for Protected Vegetable Crops (S.Y.), China Agricultural University, Beijing 100193, China.|State Key Laboratory of Plant Cell and Chromosome Engineering, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China (H.Z., L.Z., C.X., S.L., H.C.).|College of Agronomy, Southern Regional Collaborative Innovation Center for Grain and Oil Crops, Hunan Agricultural University, Changsha, Hunan 410128, China (H.L.).|Shanghai Center for Plant Stress Biology, Shanghai Institute for Biological Sciences, Chinese Academy of Sciences, Shanghai 201602, China (C.Z., R.L.); and.|University of the Chinese Academy of Sciences, Beijing 100039, China (H.Z.).
authors.forename: Hongbing
authors.initials: H
authors.lastname: Luo
authors.affiliation: National Maize Improvement Center of China, Beijing Key Laboratory of Crop Genetic Improvement, MOE Key Laboratory of Crop Heterosis and Utilization (X.C., H.S., Z.D., W.J.), and Department of Vegetable Sciences, Beijing Key Laboratory of Growth and Developmental Regulation for Protected Vegetable Crops (S.Y.), China Agricultural University, Beijing 100193, China.|State Key Laboratory of Plant Cell and Chromosome Engineering, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China (H.Z., L.Z., C.X., S.L., H.C.).|College of Agronomy, Southern Regional Collaborative Innovation Center for Grain and Oil Crops, Hunan Agricultural University, Changsha, Hunan 410128, China (H.L.).|Shanghai Center for Plant Stress Biology, Shanghai Institute for Biological Sciences, Chinese Academy of Sciences, Shanghai 201602, China (C.Z., R.L.); and.|University of the Chinese Academy of Sciences, Beijing 100039, China (H.Z.).
authors.forename: Li
authors.initials: L
authors.lastname: Zhao
authors.affiliation: National Maize Improvement Center of China, Beijing Key Laboratory of Crop Genetic Improvement, MOE Key Laboratory of Crop Heterosis and Utilization (X.C., H.S., Z.D., W.J.), and Department of Vegetable Sciences, Beijing Key Laboratory of Growth and Developmental Regulation for Protected Vegetable Crops (S.Y.), China Agricultural University, Beijing 100193, China.|State Key Laboratory of Plant Cell and Chromosome Engineering, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China (H.Z., L.Z., C.X., S.L., H.C.).|College of Agronomy, Southern Regional Collaborative Innovation Center for Grain and Oil Crops, Hunan Agricultural University, Changsha, Hunan 410128, China (H.L.).|Shanghai Center for Plant Stress Biology, Shanghai Institute for Biological Sciences, Chinese Academy of Sciences, Shanghai 201602, China (C.Z., R.L.); and.|University of the Chinese Academy of Sciences, Beijing 100039, China (H.Z.).
authors.forename: Zhaobin
authors.initials: Z
authors.lastname: Dong
authors.affiliation: National Maize Improvement Center of China, Beijing Key Laboratory of Crop Genetic Improvement, MOE Key Laboratory of Crop Heterosis and Utilization (X.C., H.S., Z.D., W.J.), and Department of Vegetable Sciences, Beijing Key Laboratory of Growth and Developmental Regulation for Protected Vegetable Crops (S.Y.), China Agricultural University, Beijing 100193, China.|State Key Laboratory of Plant Cell and Chromosome Engineering, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China (H.Z., L.Z., C.X., S.L., H.C.).|College of Agronomy, Southern Regional Collaborative Innovation Center for Grain and Oil Crops, Hunan Agricultural University, Changsha, Hunan 410128, China (H.L.).|Shanghai Center for Plant Stress Biology, Shanghai Institute for Biological Sciences, Chinese Academy of Sciences, Shanghai 201602, China (C.Z., R.L.); and.|University of the Chinese Academy of Sciences, Beijing 100039, China (H.Z.).
authors.forename: Shuangshuang
authors.initials: S
authors.lastname: Yan
authors.affiliation: National Maize Improvement Center of China, Beijing Key Laboratory of Crop Genetic Improvement, MOE Key Laboratory of Crop Heterosis and Utilization (X.C., H.S., Z.D., W.J.), and Department of Vegetable Sciences, Beijing Key Laboratory of Growth and Developmental Regulation for Protected Vegetable Crops (S.Y.), China Agricultural University, Beijing 100193, China.|State Key Laboratory of Plant Cell and Chromosome Engineering, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China (H.Z., L.Z., C.X., S.L., H.C.).|College of Agronomy, Southern Regional Collaborative Innovation Center for Grain and Oil Crops, Hunan Agricultural University, Changsha, Hunan 410128, China (H.L.).|Shanghai Center for Plant Stress Biology, Shanghai Institute for Biological Sciences, Chinese Academy of Sciences, Shanghai 201602, China (C.Z., R.L.); and.|University of the Chinese Academy of Sciences, Beijing 100039, China (H.Z.).
authors.forename: Cheng
authors.identifier: 0000-0003-0518-5924
authors.initials: C
authors.lastname: Zhao
authors.affiliation: National Maize Improvement Center of China, Beijing Key Laboratory of Crop Genetic Improvement, MOE Key Laboratory of Crop Heterosis and Utilization (X.C., H.S., Z.D., W.J.), and Department of Vegetable Sciences, Beijing Key Laboratory of Growth and Developmental Regulation for Protected Vegetable Crops (S.Y.), China Agricultural University, Beijing 100193, China.|State Key Laboratory of Plant Cell and Chromosome Engineering, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China (H.Z., L.Z., C.X., S.L., H.C.).|College of Agronomy, Southern Regional Collaborative Innovation Center for Grain and Oil Crops, Hunan Agricultural University, Changsha, Hunan 410128, China (H.L.).|Shanghai Center for Plant Stress Biology, Shanghai Institute for Biological Sciences, Chinese Academy of Sciences, Shanghai 201602, China (C.Z., R.L.); and.|University of the Chinese Academy of Sciences, Beijing 100039, China (H.Z.).
authors.forename: Renyi
authors.initials: R
authors.lastname: Liu
authors.affiliation: National Maize Improvement Center of China, Beijing Key Laboratory of Crop Genetic Improvement, MOE Key Laboratory of Crop Heterosis and Utilization (X.C., H.S., Z.D., W.J.), and Department of Vegetable Sciences, Beijing Key Laboratory of Growth and Developmental Regulation for Protected Vegetable Crops (S.Y.), China Agricultural University, Beijing 100193, China.|State Key Laboratory of Plant Cell and Chromosome Engineering, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China (H.Z., L.Z., C.X., S.L., H.C.).|College of Agronomy, Southern Regional Collaborative Innovation Center for Grain and Oil Crops, Hunan Agricultural University, Changsha, Hunan 410128, China (H.L.).|Shanghai Center for Plant Stress Biology, Shanghai Institute for Biological Sciences, Chinese Academy of Sciences, Shanghai 201602, China (C.Z., R.L.); and.|University of the Chinese Academy of Sciences, Beijing 100039, China (H.Z.).
authors.forename: Chunyan
authors.initials: C
authors.lastname: Xu
authors.affiliation: National Maize Improvement Center of China, Beijing Key Laboratory of Crop Genetic Improvement, MOE Key Laboratory of Crop Heterosis and Utilization (X.C., H.S., Z.D., W.J.), and Department of Vegetable Sciences, Beijing Key Laboratory of Growth and Developmental Regulation for Protected Vegetable Crops (S.Y.), China Agricultural University, Beijing 100193, China.|State Key Laboratory of Plant Cell and Chromosome Engineering, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China (H.Z., L.Z., C.X., S.L., H.C.).|College of Agronomy, Southern Regional Collaborative Innovation Center for Grain and Oil Crops, Hunan Agricultural University, Changsha, Hunan 410128, China (H.L.).|Shanghai Center for Plant Stress Biology, Shanghai Institute for Biological Sciences, Chinese Academy of Sciences, Shanghai 201602, China (C.Z., R.L.); and.|University of the Chinese Academy of Sciences, Beijing 100039, China (H.Z.).
authors.forename: Song
authors.initials: S
authors.lastname: Li
authors.affiliation: National Maize Improvement Center of China, Beijing Key Laboratory of Crop Genetic Improvement, MOE Key Laboratory of Crop Heterosis and Utilization (X.C., H.S., Z.D., W.J.), and Department of Vegetable Sciences, Beijing Key Laboratory of Growth and Developmental Regulation for Protected Vegetable Crops (S.Y.), China Agricultural University, Beijing 100193, China; hbchen@genetics.ac.cn weiweijin@cau.edu.cn.|State Key Laboratory of Plant Cell and Chromosome Engineering, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China (H.Z., L.Z., C.X., S.L., H.C.); hbchen@genetics.ac.cn weiweijin@cau.edu.cn.|College of Agronomy, Southern Regional Collaborative Innovation Center for Grain and Oil Crops, Hunan Agricultural University, Changsha, Hunan 410128, China (H.L.); hbchen@genetics.ac.cn weiweijin@cau.edu.cn.|Shanghai Center for Plant Stress Biology, Shanghai Institute for Biological Sciences, Chinese Academy of Sciences, Shanghai 201602, China (C.Z., R.L.); and hbchen@genetics.ac.cn weiweijin@cau.edu.cn.|University of the Chinese Academy of Sciences, Beijing 100039, China (H.Z.) hbchen@genetics.ac.cn weiweijin@cau.edu.cn.
authors.forename: Huabang
authors.initials: H
authors.lastname: Chen
authors.affiliation: National Maize Improvement Center of China, Beijing Key Laboratory of Crop Genetic Improvement, MOE Key Laboratory of Crop Heterosis and Utilization (X.C., H.S., Z.D., W.J.), and Department of Vegetable Sciences, Beijing Key Laboratory of Growth and Developmental Regulation for Protected Vegetable Crops (S.Y.), China Agricultural University, Beijing 100193, China; hbchen@genetics.ac.cn weiweijin@cau.edu.cn.|State Key Laboratory of Plant Cell and Chromosome Engineering, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China (H.Z., L.Z., C.X., S.L., H.C.); hbchen@genetics.ac.cn weiweijin@cau.edu.cn.|College of Agronomy, Southern Regional Collaborative Innovation Center for Grain and Oil Crops, Hunan Agricultural University, Changsha, Hunan 410128, China (H.L.); hbchen@genetics.ac.cn weiweijin@cau.edu.cn.|Shanghai Center for Plant Stress Biology, Shanghai Institute for Biological Sciences, Chinese Academy of Sciences, Shanghai 201602, China (C.Z., R.L.); and hbchen@genetics.ac.cn weiweijin@cau.edu.cn.|University of the Chinese Academy of Sciences, Beijing 100039, China (H.Z.) hbchen@genetics.ac.cn weiweijin@cau.edu.cn.
authors.forename: Weiwei
authors.identifier: 0000-0001-9320-9628
authors.initials: W
authors.lastname: Jin
mesh_terms: D017360:Arabidopsis / Q000235:genetics; D003001:Cloning, Molecular; D017124:Conserved Sequence / Q000235:genetics; D004269:DNA, Bacterial; D004721:Endoplasmic Reticulum / Q000378:metabolism; D005227:Fatty Acids / Q000378:metabolism; D020869:Gene Expression Profiling; D018506:Gene Expression Regulation, Plant; D017343:Genes, Plant; D008563:Membrane Lipids / Q000378:metabolism; D008954:Models, Biological; D016254:Mutagenesis, Insertional / Q000235:genetics; D009154:Mutation / Q000235:genetics; D010641:Phenotype; D019441:Plant Epidermis / Q000378:metabolism*; D010940:Plant Proteins / Q000378:metabolism*; D011058:Pollen / Q000254:growth & development* / Q000378:metabolism* / Q000648:ultrastructure; D012689:Sequence Homology, Nucleic Acid; D013347:Subcellular Fractions / Q000378:metabolism; D014885:Waxes / Q000378:metabolism; D003313:Zea mays / Q000235:genetics / Q000254:growth & development* / Q000378:metabolism* / Q000648:ultrastructure
chemical_list: D004269:DNA, Bacterial; D005227:Fatty Acids; D008563:Membrane Lipids; D010940:Plant Proteins; C036483:T-DNA; D014885:Waxes; C000521:cutin