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Plant Physiology 140:1151-1168 (2006) © 2006 American Society of Plant Biologists Integrating Membrane Transport with Male Gametophyte Development and Function through Transcriptomics1,[W]Department of Cell Biology and Molecular Genetics, University of Maryland, College Park, Maryland 20742–5815 (K.W.B., S.P., E.P.N., H.S.); Laboratory of Pollen Biology, Institute of Experimental Botany, Academy of Sciences of the Czech Republic, 16502 Prague 6, Czech Republic (D.H.); Department of Plant Physiology, Charles University, 128 44 Praha 2, Czech Republic (D.H.); Department Plant Biology, University of Minnesota, St. Paul, Minnesota 55108 (J.M.W.); Department of Pediatrics and Department of Human and Molecular Genetics, Children's Nutrition Research Center, Baylor College of Medicine, Houston, Texas 77030 (K.D.H.); and Department of Biology, University of Leicester, Leicester LE1 7RH, United Kingdom (D.T.)
Male fertility depends on the proper development of the male gametophyte, successful pollen germination, tube growth, and delivery of the sperm cells to the ovule. Previous studies have shown that nutrients like boron, and ion gradients or currents of Ca2+, H+, and K+ are critical for pollen tube growth. However, the molecular identities of transporters mediating these fluxes are mostly unknown. As a first step to integrate transport with pollen development and function, a genome-wide analysis of transporter genes expressed in the male gametophyte at four developmental stages was conducted. Approximately 1,269 genes encoding classified transporters were collected from the Arabidopsis (Arabidopsis thaliana) genome. Of 757 transporter genes expressed in pollen, 16% or 124 genes, including AHA6, CNGC18, TIP1.3, and CHX08, are specifically or preferentially expressed relative to sporophytic tissues. Some genes are highly expressed in microspores and bicellular pollen (COPT3, STP2, OPT9), while others are activated only in tricellular or mature pollen (STP11, LHT7). Analyses of entire gene families showed that a subset of genes, including those expressed in sporophytic tissues, was developmentally regulated during pollen maturation. Early and late expression patterns revealed by transcriptome analysis are supported by promoter::
1 This work was supported in part by the National Science Foundation (Arabidopsis 2010 grant nos. 0209792 to J.M.W., 020977 to K.D.H., and IBN0209788 to H.S.) and by the U.S. Department of Energy (grant no. DEFG0295ER20200 to H.S.). D.T. and D.H. gratefully acknowledge support from the Royal Society Joint Project (grant no. 2004/R3–EU). D.H. was supported through a Grant Agency of the Academy of Sciences of the Czech Republic (grant no. KJB6038409). D.T. gratefully acknowledges support from the Biotechnology and Biological Sciences Research Council and the GARNet transcriptome center at the Nottingham Arabidopsis Stock Centre for performing pollen microarray hybridizations. 2 Present address: Center for Genome Sciences, Washington University School of Medicine, Campus Box 8510, 4444 Forest Park Blvd., St. Louis, MO 63108. The author responsible for distribution of materials integral to the findings presented in this article in accordance with the policy described in the Instructions for Authors (www.plantphysiol.org) is: Heven Sze (hsze{at}umd.edu). [W] The online version of this article contains Web-only data. www.plantphysiol.org/cgi/doi/10.1104/pp.105.074708. * Corresponding author; e-mail hsze{at}umd.edu; fax 301–314–0981. Received December 1, 2005; returned for revision December 1, 2005; accepted January 13, 2006. Related articles in Plant Physiol.:
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-glucuronidase analyses of CHX genes and by other methods. Recent genetic studies based on a few transporters, including plasma membrane H+ pump AHA3, Ca2+ pump ACA9, and K+ channel SPIK, further support the expression patterns and the inferred functions revealed by our analyses. Thus, revealing the distinct expression patterns of specific transporters and unknown polytopic proteins during microgametogenesis provides new insights for strategic mutant analyses necessary to integrate the roles of transporters and potential receptors with male gametophyte development. 
