title: CsNIP2;1 is a Plasma Membrane Transporter from Cucumis sativus that Facilitates Urea Uptake When Expressed in Saccharomyces cerevisiae and Arabidopsis thaliana.
abstract: Urea is an important source of nitrogen (N) for the growth and development of plants. It occurs naturally in soils, is the major N source in agricultural fertilizers and is an important N metabolite in plants. Therefore, the identification and characterization of urea transporters in higher plants is important for the fundamental understanding of urea-based N nutrition in plants and for designing novel strategies for improving the N-use efficiency of urea based-fertilizers. Progress in this area, however, is hampered due to scarce knowledge of plant urea transporters. From what is known, urea uptake from the soil into plant roots is mediated by two types of transporters: the major intrinsic proteins (MIPs) and the DUR3 orthologs, mediating low- and high-affinity urea transport, respectively. Here we characterized a MIP family member from Cucumis sativus, CsNIP2;1, with regard to its contribution to urea transport. We show that CsNIP2;1 is a plasma membrane transporter that mediates pH-dependent urea uptake when expressed in yeast. We also found that ectopic expression of CsNIP2;1 improves growth of wild-type Arabidopsis thaliana and rescues growth and development of the atdur3-3 mutant on medium with urea as the sole N source. In addition, CsNIP2;1 is transcriptionally up-regulated by N deficiency, urea and NO3 (-). These data and results from the analyses of the pattern of CsNIP2;1 expression in A. thaliana and cucumber suggest that CsNIP2;1 might be involved in multiple steps of urea-based N nutrition, including urea uptake and internal transport during N remobilization throughout seed germination and N delivery to developing tissues.
authors: Research Center of Organic Agriculture Technology, College of Agriculture and Biotechnology, China Agricultural University, Beijing, PR China These authors contributed equally to this work. duxge@cau.edu.cn okv2@cornell.edu.
authors: Lu
authors: L
authors: Zhang
authors: School of Integrative Plant Sciences, Soil and Crop Sciences Section, Cornell University, Ithaca, NY, USA These authors contributed equally to this work. duxge@cau.edu.cn okv2@cornell.edu.
authors: Jiapei
authors: J
authors: Yan
authors: School of Integrative Plant Sciences, Soil and Crop Sciences Section, Cornell University, Ithaca, NY, USA duxge@cau.edu.cn okv2@cornell.edu.
authors: Olena K
authors: OK
authors: Vatamaniuk
authors: Research Center of Organic Agriculture Technology, College of Agriculture and Biotechnology, China Agricultural University, Beijing, PR China duxge@cau.edu.cn okv2@cornell.edu.
authors: Xiangge
authors: X
authors: Du
mesh_terms: D000595:Amino Acid Sequence; D017360:Arabidopsis / Q000187:drug effects / Q000235:genetics* / Q000254:growth & development; D002462:Cell Membrane / Q000187:drug effects / Q000378:metabolism*; D018553:Cucumis sativus / Q000187:drug effects / Q000378:metabolism*; D018506:Gene Expression Regulation, Plant / Q000187:drug effects; D005816:Genetic Complementation Test; D005990:Glycerol / Q000494:pharmacology; D026901:Membrane Transport Proteins / Q000737:chemistry / Q000378:metabolism*; D009154:Mutation / Q000235:genetics; D009566:Nitrates / Q000378:metabolism; D009584:Nitrogen / Q000494:pharmacology; D010940:Plant Proteins / Q000737:chemistry / Q000378:metabolism*; D018517:Plant Roots / Q000187:drug effects / Q000378:metabolism; D030821:Plants, Genetically Modified; D021381:Protein Transport / Q000187:drug effects; D011523:Protoplasts / Q000187:drug effects / Q000378:metabolism; D012441:Saccharomyces cerevisiae / Q000378:metabolism*; D014170:Transformation, Genetic / Q000187:drug effects; D015854:Up-Regulation / Q000187:drug effects; D014508:Urea / Q000378:metabolism*
chemical_list: D026901:Membrane Transport Proteins; D009566:Nitrates; D010940:Plant Proteins; D014508:Urea; D009584:Nitrogen; D005990:Glycerol
keywords: CsNIP2;1; Cucumis sativus; MIP; NIP; Protoplasts; Urea transport