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Jiahua Xie, Todd C. Wehner, Kurt Wollenberg, Michael D. Purugganan, and Mark A. Conkling

Aquaporin proteins are part of an ancient family that functions as water transporting facilitators in all organisms. Phylogenetic and physiological analyses have revealed that plant aquaporins consist of two groups: the plasma membrane intrinsic proteins (PIPs) and the tonoplast intrinsic proteins (TIPs). Using the conserved asparagineproline-alanine (NPA) to NPA motif regions, we studied the evolution of 35 plant aquaporins that included nine of our newly cloned cucumber aquaporins and 26 from the GenBank database. Results indicated that NPA repeated regions were effective for phylogenetically characterizing the plant aquaporin family, and to accurately localize the introns. Phylogenetic analysis showed that 35 plant aquaporins fell into two distinct groups (except for the Arabidopsis gene AtMip)—PIPs and TIPs. The nine cucumber aquaporins belong to the PIP group that were localized further into two different sub-groups. The intron analysis showed that introns of plant aquaporins mainly consist of two types. Eighteen PIPs shared identical intron positions localized in connecting loop C between amino acids 95 and 96. Nine TIPs shared the other identical intron positions localized in connecting loop D between amino acids 44 and 45. Cucumber aquaporins CRB9 and CRB10 (with no intron in the repeated NPA regions) may be the result of intron loss events, while intronless rice (Orzya sativa) Os-TIP1 and Os-TIP2 may have resulted from other intron loss events. PIP11 and Os-PIP do not have the same amino acid number as major PIP members, but combined phylogenetic analysis results along with intron positions and phases showed that they belong to the PIP group. The phylogenetic tree and intron position information suggest that AtMip was mis-annotated as a member of aquaporin, and is a homologue of the glycerol facilitator-like protein. Introns share identical positions and phases within the PIP group (except PIP13) or the TIP group, but differ between the plasma and the tonoplast membrane aquaporins matching the phylogenetic analysis results. Intron positions of the repeated NPA regions of plant aquaporins that have stable inheritance can act as molecular markers for phylogenetic studies.

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Lili Zhuang, Mengxian Liu, Xiuyun Yuan, Zhimin Yang, and Bingru Huang

aquaporin genes in Nicotiana tabacum plants Plant Cell Physiol. 49 801 813 Maurel, C. 2007 Plant aquaporins: Novel functions and regulation properties FEBS Lett. 581 2227 2236 Maurel, C. Verdoucq, L. Luu, D. Santoni, V. 2008 Plant aquaporins: Membrane

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W. Roland Leatherwood, D. Mason Pharr, Lisa O. Dean, and John D. Williamson

, M.J. Mas, A. 2002 Plant aquaporins Physiol. Plant. 115 175 182 Bray, E.A. Bailey-Serres, J. Weretilnyk, E. 2000 Responses to abiotic stresses 1158 1203 Buchanan B

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Daniel Leskovar and Yahia Othman

alter growth of vegetable transplants HortTechnology 22 56 63 Maurel, C. Verdoucq, L. Luu, D. Santoni, V. 2008 Plant aquaporins: Membrane channels with multiple integrated functions Annu. Rev. Plant Biol. 59 595 624 McDonald, J. Ericsson, T. Larsson, C

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César Mota-Cadenas, Carlos Alcaraz-López, M. Carmen Martínez-Ballesta, and Micaela Carvajal

nutrition of higher plants 2nd Ed Academia Press London, UK Martínez-Ballesta, M.C. Silva, C. López-Berenguer, C. Cabañero, F.J. Carvajal, M. 2006 Plant aquaporins: New perspectives on water and nutrient uptake in saline environment Plant Biol. 8 535 546

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Manuel G. Astacio and Marc W. van Iersel

expansion Plant Physiol. 143 1905 1917 Tyerman, S.D. Niemietz, C.M. Bramley, H. 2002 Plant aquaporins: Multifunctional water and solute channels with expanding roles Plant Cell Environ. 25 173 194 van Iersel, M.W. Seader, K. Dove, S. 2009 Exogenous abscisic

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Xiaobo Sun, Yanming Deng, Lijian Liang, Xinping Jia, Zheng Xiao, and Jiale Su

, X. Yang, Y. 2014 Molecular cloning and expression analysis of tea plant aquaporin (AQP) gene family Plant Physiol. Biochem. 83 65 76 Zelazny, E. Borst, J.W. Muylaert, M. Batoko, H. Hemminga, M.A. Chaumont, F. 2007 FRET imaging in living maize cells

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Kaori Ando and Rebecca Grumet

resistance and fruit quality traits in melon ( Cucumis melo L.) BMC Genomics 10 467 Maurel, C. Verdoucq, L. Luu, D.-T. Santoni, V. 2008 Plant aquaporins: Membrane channels with multiple integrated