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Research ArticleGENOME ANALYSIS
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The Complete Set of Genes Encoding Major Intrinsic Proteins in Arabidopsis Provides a Framework for a New Nomenclature for Major Intrinsic Proteins in Plants

Urban Johanson, Maria Karlsson, Ingela Johansson, Sofia Gustavsson, Sara Sjövall, Laure Fraysse, Alfons R. Weig, Per Kjellbom
Urban Johanson
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Maria Karlsson
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Ingela Johansson
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Sofia Gustavsson
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Sara Sjövall
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Laure Fraysse
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Alfons R. Weig
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Per Kjellbom
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Published August 2001. DOI: https://doi.org/10.1104/pp.126.4.1358

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    Fig. 1.

    Phylogenetic comparison of the complete set of 35 different MIPs encoded in the Arabidopsis genome. Plant MIPs are divided into four distinct subfamilies: PIPs, TIPs, NIPs, and SIPs. Similar proteins within a subfamily, with a maximum of 30% divergence, are clustered in monophyletic groups. The first and the last digit in the protein name identify the group and the individual gene product, respectively. This tree was obtained using the whole alignment and the distance method. Omitting the none-conserved N- and C-terminal regions from the phylogenetic analysis does not break the defined groups; only the relative positions of PIP2;6 and TIP3s is changed. In this case PIP2;6 forms a separate branch between PIP2;4 and PIP2;5 and TIP3s branch between TIP2s andTIP4;1. The bar indicates the mean distance of 0.05 changes per amino acid residue.

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    Fig. 2.

    Bootstrap majority-rule consensus tree generated with the distance method. A bootstrap value of 100% indicates branches that were supported in all replicates of resampling of data. Branches with a bootstrap value of less than 50% are collapsed.

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    Fig. 3.

    Schematic structure of MIP encoding genes in Arabidopsis. Horizontal bars and gaps depict exons and intron positions, respectively. Parts encoding transmembrane helices H1 to H6 according to an alignment with GlpF are indicated by vertical bars. The color on the vertical bars shows homologous transmembrane helices in the first and second halves of the MIPs. The exons and transmembrane helices are drawn to scale but the positions of helices are schematic. Helices encoded on two exons are only indicated on the exon where the major part is encoded. Small indels in the alignment of different genes, positioned between two helices on the same exon, are not shown.

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    Table I.

    Proposed new names for all Arabidopsis MIPs

    New Name1-aJohanson et al.1-bWeig et al.1-cAGI Gene1-dBAC Gene1-eAccession No.1-fExpressed Sequence Tag (EST)1-gComments1-h
    PIP1;1PIP1aPIP1aAt3g61430F2A19.30 CAB71073 Yes
    PIP1;2PIP1bPIP1bAt2g45960F4I18.6 AAC28529 YesAthH2, TMP-A
    PIP1;3PIP1cPIP1cAt1g01620F22L4.16 AAF81320 YesTMP-B
    PIP1;4PIP1eTMP-CAt4g00430F5I10.2 AAF02782 YesExon 2 ends at 86,121, not 86,097; compare with TMP-C.
    PIP1;5PIP1dpCR55At4g23400F16G20.100 CAA20461 Yes
    PIP2;1PIP2aPIP2aAt3g53420F4P12_120 CAB67649 Yes
    PIP2;2PIP2bPIP2bAt2g37170T2N18.7 AAD18142 Yes
    PIP2;3PIP2cRD28At2g37180T2N18.6 AAD18141 Yes
    PIP2;4PIP2f T04164 At5g60660MUP24.8 BAB09839 Yes
    PIP2;5PIP2d–At3g54820F28P10.200 CAB41102 Yes
    PIP2;6PIP2e T22419 At2g39010T7F6.18 AAC79629 Yes
    PIP2;7PIP3aPIP3At4g35100M4E13.150 CAA17774 YesSalt-stress-inducible MIP
    PIP2;8PIP3b–At2g16850F12A24.3 AAC64216 Yes
    –Pseudo PIP3–At2g16830F12A24.1 AAC64228 ?Pseudogene encoding 26 amino acids with second NPA. Similar to F12A24.3.
    ––––T8O111-i 77,410–77,455?Partial F5I10.2-like sequence encoding helix 4.
    ––––F5I101-i 77,564–77,656?Partial F5I10.2-like sequence encoding loop B and helix 3.
    TIP1;1γTIP1γTIPAt2g36830T1J8.1AAD31569Yes
    TIP1;2γTIP2 Z30833, T21060,T22237 At3g26520MFE16.3 BAB01832 YesSalt-stress-inducible TIP, γTIP2
    TIP1;3γTIP3–At4g01470F11O4.1 AAC62778 ?No introns
    TIP2;1δTIP1δTIPAt3g16240MYA6.5 BAB01264 Yes
    TIP2;2δTIP2 Z18142 At4g17340d14705w CAB10515 Yes
    TIP2;3δTIP3 T20432,T76151 At5g47450MNJ7.4 BAB09071 Yes
    TIP3;1αTIPαTIPAt1g73190T18K17.14 AAG52132 Yes
    TIP3;2βTIPβTIPAt1g17810F2H15.4 AAF97261 Yes
    TIP4;1ɛTIP T21742 At2g25810F17H15.16 AAC42249 YesSimilar to NtTIPa
    TIP5;1ζTIP–At3g47440T21L8.190 CAB51216 ?
    –Pseudo-γTIP2––F7P11-i 79,395–79,728?Pseudogene similar to MFE16.3, truncated, NPV instead of NPA, no exon 2.
    –Pseudo-δTIP–At1g52180F9I5.3 AAF29403 ?Pseudogene, only exon 2, 124 amino acids.
    NIP1;1NLM1NLM1At4g19030F13C5.200 CAA16760 YesMisannotated: delete amino acids 205 through 216; exon 4 starts at 86,879, not 86,915.
    NIP1;2NLM2NLM2At4g18910F13C5.80 CAA16748 Yes
    NIP2;1NLM3NLM4At2g34390T31E10.27 AAC26712 Yes1-j
    NIP3;1NLM9–At1g318801-k F5M6.28 AAG50717 ?F5M6.28 is missing exon 1; exon 1, 38,681 through 38,550; exon 2 starts at 38,310, long C terminus.
    NIP4;1NLM4–At5g37810K22F20.9 BAB10360 ?
    NIP4;2NLM5NLM5At5g37820K22F20.10 BAB10361 Yes1-j
    NIP5;1NLM6NLM8At4g10380F24G24.180 CAB39791 YesLong N terminus.
    NIP6;1NLM7NLM7At1g80760F23A5.11 AAF14664 YesLong N terminus.
    NIP7;1NLM8NLM6At3g06100F28L1.3 AAF30303 YesIntron 3, 8,151 through 7,945, avoids TT acceptor splice site, restores conserved length of exon 4.
    –Pseudo-NLM3NLM3At2g29870T27A16.3 AAC35214 ?Pseudogene encoding 139 amino acids, missing first NPA.
    –Pseudo-NLM9–At2g21020F26H11.22 AAD29814 ?Pseudogene, version 3 missing second NPA. Alternate annotation results in two NPAs, but missing helix 3 and 4, long C terminus.
    SIP1;1SIP1a–At3g04090T6K12.29 AAF26804 Yes
    SIP1;2SIP1b–At5g18290MRG7.25 BAB09487 Yes
    SIP2;1SIP2–At3g56950F24I3.30 CAB72165 Yes
    • ↵F1-a  Proposed new names for Arabidopsis MIPs. When referring to the corresponding gene, the name is written in italics. Genes encoding partial MIP-like sequences without all transmembrane regions are regarded as nonfunctional pseudogenes. Pseudogenes are not included in the proposed nomenclature as indicated by “–”.

    • ↵F1-b  Nomenclature used inJohansson et al. (2000) and in U. Johanson and S. Gustavsson (unpublished data).

    • ↵F1-c  Nomenclature used in Weig et al. (1997) and Weig and Jakob (2000a, 2000b).

    • ↵F1-d  Arabidopsis Genome Initiative (AGI) gene nomenclature; At3g56950 indicates gene no. 5,695 (counting from top) on chromosome 3.

    • ↵F1-e  Gene name used in annotation of bacterial artificial chromosome (BAC) or P1 sequences.

    • ↵F1-f  Protein accession no. for a translation of the corresponding gene. When a gene annotation is missing, the location in the BAC is given in base pairs.

    • ↵F1-g  Matching EST found in databases. ?, Not found.

    • ↵F1-h  Alternative exon/introns positions proposed and used in this paper and odd features of the gene or the encoded protein. Some older names found in the literature (Kaldenhoff et al., 1993; Shagan and Bar-Zvi, 1993; Shagan et al., 1993; Pih et al., 1999) and in Genbank accession no.AF057137.

    • ↵F1-i  Not annotated.

    • ↵F1-j  cDNA cloned by reverse transcriptase-PCR (Weig and Jakob, 2000b).

    • ↵F1-k  At1g31880 has 285 amino acids added at the N terminus, previously annotated as a separate gene (F5M6.11). At1g31880 is missing 29 amino acids close to the C terminus.

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The Complete Set of Genes Encoding Major Intrinsic Proteins in Arabidopsis Provides a Framework for a New Nomenclature for Major Intrinsic Proteins in Plants
Urban Johanson, Maria Karlsson, Ingela Johansson, Sofia Gustavsson, Sara Sjövall, Laure Fraysse, Alfons R. Weig, Per Kjellbom
Plant Physiology Aug 2001, 126 (4) 1358-1369; DOI: 10.1104/pp.126.4.1358

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The Complete Set of Genes Encoding Major Intrinsic Proteins in Arabidopsis Provides a Framework for a New Nomenclature for Major Intrinsic Proteins in Plants
Urban Johanson, Maria Karlsson, Ingela Johansson, Sofia Gustavsson, Sara Sjövall, Laure Fraysse, Alfons R. Weig, Per Kjellbom
Plant Physiology Aug 2001, 126 (4) 1358-1369; DOI: 10.1104/pp.126.4.1358
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Plant Physiology: 126 (4)
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