TOR介导的糖激活SAM活性的拟南芥自然变异位点鉴定
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摘要
在黑暗条件下,糖能激活拟南芥的茎顶端分生组织(SAM)发育成叶状器官,该现象已成为植物糖敏感性的重要指标之一。本研究发现,通过RNAi降低雷帕霉素靶标(TOR)的表达,能降低拟南芥幼苗的糖敏感性,表明TOR参与了糖对SAM的激活。本研究还通过对265个拟南芥自然生态型的鉴定,筛选到了一个糖敏感性较低的生态型CS76205;进一步分析发现, Col-0和CS76205的F_1代的糖敏感性处于Col-0和CS76205之间,偏向CS76205; Col-0×CS76205 F_2群体的糖敏感性呈正态分布,通过二代测序对CS76205生态型和F_2代的两个极端表型混合池进行全基因组重测序,在3号染色体定位到一个糖激活SAM活性相关的区域,位于2.520 Mb和3.715Mb之间,利用生物信息学的手段对该关联区域进行注释,筛选出9个关键的候选基因。这为进一步克隆该位点基因、揭示该基因功能及其和TOR之间的关系奠定基础。
In the dark, sugar activates the shoot apical meristem(SAM) of Arabidopsis thaliana to develop leaflike organs, which has become one of the important indicators of sugar sensitivity in plant. Our study found that the RNAi of target of rapamycin(TOR) decreased the sugar sensitivity, indicating that TOR is involved in the sugar-responsive SAM activation. We obtained a sugar less sensitive ecotype, CS76205, from 265 Arabidopsis natural ecotypes. Further analysis showed that the degree of sugar sensitivity of F_1 of Col-0 × CS76205 was intermediate between Col-0 and CS76205 and biased towards CS76205. The sugar sensitivity of Col-0 ×CS76205 F_2 population was normal distribution. Whole-genome resequencing and subsequent Bulk Segregant Analysis(BSA)-seq analysis suggested that the sugar-induced SAM activity was mapped on chromosome 3 through a next-generation sequencing of CS76205 and the two extreme phenotype pools of F_2 generation. The associated region, located between 2.520 Mb and 3.715 Mb, was annotated with bioinformatics to filter out nine key candidate genes. Our study will contributes to lay the foundation for further cloning of the locus gene, revealing the function of the gene and its relationship with TOR.
In the dark, sugar activates the shoot apical meristem(SAM) of Arabidopsis thaliana to develop leaflike organs, which has become one of the important indicators of sugar sensitivity in plant. Our study found that the RNAi of target of rapamycin(TOR) decreased the sugar sensitivity, indicating that TOR is involved in the sugar-responsive SAM activation. We obtained a sugar less sensitive ecotype, CS76205, from 265 Arabidopsis natural ecotypes. Further analysis showed that the degree of sugar sensitivity of F_1 of Col-0 × CS76205 was intermediate between Col-0 and CS76205 and biased towards CS76205. The sugar sensitivity of Col-0 ×CS76205 F_2 population was normal distribution. Whole-genome resequencing and subsequent Bulk Segregant Analysis(BSA)-seq analysis suggested that the sugar-induced SAM activity was mapped on chromosome 3 through a next-generation sequencing of CS76205 and the two extreme phenotype pools of F_2 generation. The associated region, located between 2.520 Mb and 3.715 Mb, was annotated with bioinformatics to filter out nine key candidate genes. Our study will contributes to lay the foundation for further cloning of the locus gene, revealing the function of the gene and its relationship with TOR.
引文
Anderson GH,Veit B,Hanson MR(2005).The Arabidopsis AtRaptor genes are essential for post-embryonic plant growth.BMC Biol,3:12
Bach L,Faure JD(2010).Role of very-long-chain fatty acids in plant development,when chain length does matter.C RBiol,333(4):361-370
Baena-Gonzalez E,Rolland F,Thevelein JM,et al(2007).Acentral integrator of transcription networks in plant stress and energy signalling.Nature,448(7156):938-942
Baena-Gonzalez E,Sheen J(2008).Convergent energy and stress signaling.Trends Plant Sci,13(9):474-482
Baier M,Hemmann G,Holman R,et al(2004).Characterization of mutants in Arabidopsis showing increased sugar-specific gene expression,growth,and developmental responses.Plant Physiol,134(1):81-91
Barton MK(2010).Twenty years on:the inner workings of the shoot apical meristem,a developmental dynamo.Dev Biol,341(1):95-113
Chaiwanon J,Wang W,Zhu JY,et al(2016).Information Integration and Communication in Plant Growth Regulation.Cell,164(6):1257-1268
Chen K,Wallis JW,McLellan MD,et al(2009).BreakDancer:an algorithm for high resolution mapping of genomic structural variation.Nat Methods,6(9):677-681
Cingolani P,Platts A,Coon M,et al(2012).A program for annotating and predicting the effects of single nucleotide polymorphisms,SnpEff:SNPs in the genome of Drosophila melanogaster strain w1118;iso-2;iso-3.Fly,6(2):80-92
Costaglioli P,Joubes J,Garcia C(2005).Profiling candidate genes involved in wax biosynthesis in Arabidopsis thaliana by microarray analysis.Biochim Biophys Acta,1734(3):247-258
De Veylder L,Segers G,Glab N,et al(1997).The Arabidopsis Cks1At protein binds the cyclin-dependent kinases Cdc2aAt and Cdc2bAt.FEBS Lett,412(3):446-452
Dunn TM,Lynch DV,Michaelson LV,et al(2004).A post-genomic approach to understanding sphingolipid metabolism in Arabidopsis thaliana.Ann Bot,93(5):483-497
Francis D,Halford NG(2006).Nutrient sensing in plant meristems.Plant Mol Biol,60:981-993
Gomez LD,Gilday A,Feil R,et al(2010).AtTPS1-mediated trehalose 6-phosphate synthesis is essential for embryogenic and vegetative growth and responsiveness to ABA in germinating seeds and stomatal guard cells.Plant J,64(1):1-13
Hanson J,Hanssen M,WieseA,et al(2008).The sucrose regulated transcription factor bZIP11 affects amino acid metabolism by regulating the expression of ASPARAGINESYNTHETASE1 and PROLINE DEHYDROGENASE2.Plant J,53(6):935-949
Hardie DG(2007).AMPK and SNF1:snuffing out stress.Cell Metab,6(5):339-340
Harris MA,Clark J,Ireland A,et al(2004).The Gene Ontology(GO)database and informatics resource.Nucleic Acids Res,32:D258-D261
Haslam TM,Kunst L(2013).Extending the story of verylong-chain fatty acid elongation.Plant Sci,210:93-107
Jensen LJ,Julien P,Kuhn M,et al(2008).eggNOG:automated construction and annotation of orthologous groups of genes.Nucleic Acids Res,36:250-254
Jeon BW,Hwang JU,Hwang Y,et al(2008).The Arabidopsis small G protein ROP2 is activated by light in guard cells and inhibits light-induced stomatal opening.Plant Cell,20(1):75-87
Joubes J,Raffaele S,Bourdenx B,et al(2008).The VLCFAelongase gene family in Arabidopsis thaliana:phylogenetic analysis,3D modelling and expression profiling.Plant Mol Biol,67(5):547-566
Kanehisa M,Goto S(2000).KEGG:kyoto encyclopedia of genes and genomes.Nucleic Acids Res,28(1):27-30
Kimura Y,Takeoka Y,Inoue M,et al(2011).Double-knockout of putative endo-β-N-acetylglucosaminidase(ENGase)genes in Arabidopsis thaliana:loss of ENGase activity induced accumulation of high-mannose type free N-glycans bearing N,N′-acetylchitobiosyl unit.Biosci Biotechnol Biochem,75(5):1019-1021
Lastdrager J,Hanson J,Smeekens S(2014).Sugar signals and the control of plant growth and development.J Exp Bot,65(3):799-807
Li H,Durbin R(2009).Fast and accurate short read alignment with Burrows-Wheeler transform.Bioinformatics,25(14):1754-1760
Li H,Handsaker B,Wysoker A,et al(2009).The Sequence Alignment/Map format and SAMtools.Bioinformatics,25(16):2078-2079
Li H,Shen JJ,Zheng ZL,et al(2001).The Rop GTPase switch controls multiple developmental processes in Arabidopsis.Plant Physiol,126(2):670-684
Li X,Cai W,Liu Y,et al(2017).Differential TOR activation and cell proliferation in Arabidopsis root and shoot apexes.Proc Natl Acad Sci USA,114(10):2765-2770
Li Y,Smith C,Corke F(2007).Signaling from an altered cell wall to the nucleus mediates sugar-responsive growth and development in Arabidopsis thaliana.Plant Cell,19(8):2500-2515
Mahfouz MM,Kim S,Delauney AJ(2006).Arabidopsis TAR-GET OF RAPAMYCIN interacts with RAPTOR,which regulates the activity of S6 kinase in response to osmotic stress signals.Plant Cell,18(2):477-490
Margalha L,Valerio C,Baena-Gonzalez E(2016).Plant SnRK1 Kinases:Structure,Regulation,and Function.In:Cordero MD,Viollet B(eds).AMP-activated Protein Kinase.Switzerland:Spinger,403-438
Martinez-Barajas E,Delatte T,Schluepmann H,et al(2011).Wheat grain development is characterized by remarkable trehalose 6-phosphate accumulation pregrain filling:tissue distribution and relationship to SNF1-related protein kinase1 activity.Plant Physiol,156(1):373-381
Mortensen SA,Sonderkaer M,et al(2011).Reduced expression of the DOG1 gene in Arabidopsis mutant seeds lacking the transcript elongation factor TFIIS.FEBS Lett,585(12):1929-1933
McKenna A,Hanna M,Banks E,et al(2010).The Genome Analysis Toolkit:a MapReduce framework for analyzing next-generation DNA sequencing data.Genome Res,20(9):1297-1303
Moore B,Zhou L,Rolland F,et al(2003).Role of the Arabidopsis glucose sensor HXK1 in nutrient,light,and hormonal signaling.Science,300(5617):332-336
Moreau M,Azzopardi M,Clement G,et al(2012).Mutations in the Arabidopsis homolog of LST8/GβL,a partner of the Target of Rapamycin kinase,impair plant growth,flowering,and metabolic adaptation to long days.Plant Cell,24(2):463-481
Peng Y,Chen L,Li S,et al(2018).BRI1 and BAK1 interact with G proteins and regulate sugar-responsive growth and development in Arabidopsis.Nat Commun,9(1):1522
Riou-Khamlichi C,Menges M,Healy JM,et al(2000).Sugar control of the plant cell cycle:differential regulation of Arabidopsis D-type cyclin gene expression.Mol Cell Biol,20(13):4513-4521
Robaglia C,Thomas M,Meyer C(2012).Sensing nutrient and energy status by SnRK1 and TOR kinases.Curr Opin Plant Biol,15(3):301-307
Roldan M,Gomez-Mena C,Ruiz-Garcia L,et al(1999).Sucrose availability on the aerial part of the plant promotes morphogenesis and flowering of Arabidopsis in the dark.Plant J,20(5):581-590
Schoof H,Lenhard M,Haecker A,et al(2000).The stem cell population of Arabidopsis shoot meristems in maintained by a regulatory loop between the CLAVATA and WUS-CHEL genes.Cell,100(6):635-644
Skylar A,Sung F,Hong F,et al(2011).Metabolic sugar signal promotes Arabidopsis meristematic proliferation via G2.Dev Biol,351(1):82-89
Smeekens S,Ma J,Hanson J,et al(2010).Sugar signals and molecular networks controlling plant growth.Curr Opin Plant Biol,13(3):274-279
Takagi H,Abe A,Yoshida K,et al(2013).QTL-seq:rapid mapping of quantitative trait loci in rice by whole genome resequencing of DNA from two bulked populations.Plant J,74(1):174-183
Timson D(2007).GHMP kinases--structures,mechanisms and potential for therapeutically relevant inhibition.Curr Enzym Inhib,3(1):77-94
Tsai AY,Gazzarrini S(2014).Trehalose-6-phosphate and SnRK1 kinases in plant development and signaling:the emerging picture.Front Plant Sci,5:1-11
UniProt Consortium(2015).UniProt:a hub for protein information.Nucleic Acids Res,43:204-212
Wang G,Ellendorff U,Kemp B,et al(2008).A genome-wide functional investigation into the roles of receptor-like proteins in Arabidopsis.Plant Physiol,147(2):503-517
Wang Y,Li J(2008).Molecular basis of plant architecture.Annu Rev Plant Biol,59:253-279
Wolf S,Ladwig F,Sticht C,et al(2014).A receptor-like protein mediates the response to pectin modification by activating brassinosteroid signaling.Proc Natl Acad Sci USA,111(42):15261-15266
Wu J,Liu Z,Zhang Z,et al(2016).Transcriptional regulation of receptor-like protein genes by environmental stresses and hormones and their overexpression activities in Arabidopsis thaliana.J Exp Bot,67(11):3339-3351
Wu X,Dabi T,Weigel D(2005).Requirement of homeobox gene STIMPY/WOX9 for Arabidopsis meristem growth and maintenance.Curr Biol,15(5):436-440
Xiong F,Zhang R,Meng Z,et al(2017).Brassinosteriod Insensitive 2(BIN2)acts as a downstream effector of the Target of Rapamycin(TOR)signaling pathway to regulate photoautotrophic growth in Arabidopsis.New Phytol,213(1):233-249
Xiong Y,Sheen J(2012).Rapamycin and glucose-target of rapamycin(TOR)protein signaling in plants.J Biol Chem,287(4):2836-2842
Yang T,Bar-Peled L,Gebhart L,et al(2009).Identification of galacturonic acid-1-phosphate kinase,a new member of the GHMP kinase superfamily in plants,and comparison with galactose-1-phosphate kinase.J Biol Chem,284(32):21526-21535
Zhang Y,Liu Z,Wang J,et al(2015).Brassinosteroid is required for sugar promotion of hypocotyl elongation in Arabidopsis in darkness.Planta,242(4):881-893
Zhang Z,Zhu JY,Roh J,et al(2016).TOR signaling promotes accumulation of BZR1 to balance growth with carbon availability in Arabidopsis.Curr Biol,26(14):1854-1860
Zheng L,Shang L,Chen X,et al(2015).TANG1,encoding a Symplekin_C domain-contained protein,influences sugar responses in Arabidopsis.Plant Physiol,168(3):1000-1012
Bach L,Faure JD(2010).Role of very-long-chain fatty acids in plant development,when chain length does matter.C RBiol,333(4):361-370
Baena-Gonzalez E,Rolland F,Thevelein JM,et al(2007).Acentral integrator of transcription networks in plant stress and energy signalling.Nature,448(7156):938-942
Baena-Gonzalez E,Sheen J(2008).Convergent energy and stress signaling.Trends Plant Sci,13(9):474-482
Baier M,Hemmann G,Holman R,et al(2004).Characterization of mutants in Arabidopsis showing increased sugar-specific gene expression,growth,and developmental responses.Plant Physiol,134(1):81-91
Barton MK(2010).Twenty years on:the inner workings of the shoot apical meristem,a developmental dynamo.Dev Biol,341(1):95-113
Chaiwanon J,Wang W,Zhu JY,et al(2016).Information Integration and Communication in Plant Growth Regulation.Cell,164(6):1257-1268
Chen K,Wallis JW,McLellan MD,et al(2009).BreakDancer:an algorithm for high resolution mapping of genomic structural variation.Nat Methods,6(9):677-681
Cingolani P,Platts A,Coon M,et al(2012).A program for annotating and predicting the effects of single nucleotide polymorphisms,SnpEff:SNPs in the genome of Drosophila melanogaster strain w1118;iso-2;iso-3.Fly,6(2):80-92
Costaglioli P,Joubes J,Garcia C(2005).Profiling candidate genes involved in wax biosynthesis in Arabidopsis thaliana by microarray analysis.Biochim Biophys Acta,1734(3):247-258
De Veylder L,Segers G,Glab N,et al(1997).The Arabidopsis Cks1At protein binds the cyclin-dependent kinases Cdc2aAt and Cdc2bAt.FEBS Lett,412(3):446-452
Dunn TM,Lynch DV,Michaelson LV,et al(2004).A post-genomic approach to understanding sphingolipid metabolism in Arabidopsis thaliana.Ann Bot,93(5):483-497
Francis D,Halford NG(2006).Nutrient sensing in plant meristems.Plant Mol Biol,60:981-993
Gomez LD,Gilday A,Feil R,et al(2010).AtTPS1-mediated trehalose 6-phosphate synthesis is essential for embryogenic and vegetative growth and responsiveness to ABA in germinating seeds and stomatal guard cells.Plant J,64(1):1-13
Hanson J,Hanssen M,WieseA,et al(2008).The sucrose regulated transcription factor bZIP11 affects amino acid metabolism by regulating the expression of ASPARAGINESYNTHETASE1 and PROLINE DEHYDROGENASE2.Plant J,53(6):935-949
Hardie DG(2007).AMPK and SNF1:snuffing out stress.Cell Metab,6(5):339-340
Harris MA,Clark J,Ireland A,et al(2004).The Gene Ontology(GO)database and informatics resource.Nucleic Acids Res,32:D258-D261
Haslam TM,Kunst L(2013).Extending the story of verylong-chain fatty acid elongation.Plant Sci,210:93-107
Jensen LJ,Julien P,Kuhn M,et al(2008).eggNOG:automated construction and annotation of orthologous groups of genes.Nucleic Acids Res,36:250-254
Jeon BW,Hwang JU,Hwang Y,et al(2008).The Arabidopsis small G protein ROP2 is activated by light in guard cells and inhibits light-induced stomatal opening.Plant Cell,20(1):75-87
Joubes J,Raffaele S,Bourdenx B,et al(2008).The VLCFAelongase gene family in Arabidopsis thaliana:phylogenetic analysis,3D modelling and expression profiling.Plant Mol Biol,67(5):547-566
Kanehisa M,Goto S(2000).KEGG:kyoto encyclopedia of genes and genomes.Nucleic Acids Res,28(1):27-30
Kimura Y,Takeoka Y,Inoue M,et al(2011).Double-knockout of putative endo-β-N-acetylglucosaminidase(ENGase)genes in Arabidopsis thaliana:loss of ENGase activity induced accumulation of high-mannose type free N-glycans bearing N,N′-acetylchitobiosyl unit.Biosci Biotechnol Biochem,75(5):1019-1021
Lastdrager J,Hanson J,Smeekens S(2014).Sugar signals and the control of plant growth and development.J Exp Bot,65(3):799-807
Li H,Durbin R(2009).Fast and accurate short read alignment with Burrows-Wheeler transform.Bioinformatics,25(14):1754-1760
Li H,Handsaker B,Wysoker A,et al(2009).The Sequence Alignment/Map format and SAMtools.Bioinformatics,25(16):2078-2079
Li H,Shen JJ,Zheng ZL,et al(2001).The Rop GTPase switch controls multiple developmental processes in Arabidopsis.Plant Physiol,126(2):670-684
Li X,Cai W,Liu Y,et al(2017).Differential TOR activation and cell proliferation in Arabidopsis root and shoot apexes.Proc Natl Acad Sci USA,114(10):2765-2770
Li Y,Smith C,Corke F(2007).Signaling from an altered cell wall to the nucleus mediates sugar-responsive growth and development in Arabidopsis thaliana.Plant Cell,19(8):2500-2515
Mahfouz MM,Kim S,Delauney AJ(2006).Arabidopsis TAR-GET OF RAPAMYCIN interacts with RAPTOR,which regulates the activity of S6 kinase in response to osmotic stress signals.Plant Cell,18(2):477-490
Margalha L,Valerio C,Baena-Gonzalez E(2016).Plant SnRK1 Kinases:Structure,Regulation,and Function.In:Cordero MD,Viollet B(eds).AMP-activated Protein Kinase.Switzerland:Spinger,403-438
Martinez-Barajas E,Delatte T,Schluepmann H,et al(2011).Wheat grain development is characterized by remarkable trehalose 6-phosphate accumulation pregrain filling:tissue distribution and relationship to SNF1-related protein kinase1 activity.Plant Physiol,156(1):373-381
Mortensen SA,Sonderkaer M,et al(2011).Reduced expression of the DOG1 gene in Arabidopsis mutant seeds lacking the transcript elongation factor TFIIS.FEBS Lett,585(12):1929-1933
McKenna A,Hanna M,Banks E,et al(2010).The Genome Analysis Toolkit:a MapReduce framework for analyzing next-generation DNA sequencing data.Genome Res,20(9):1297-1303
Moore B,Zhou L,Rolland F,et al(2003).Role of the Arabidopsis glucose sensor HXK1 in nutrient,light,and hormonal signaling.Science,300(5617):332-336
Moreau M,Azzopardi M,Clement G,et al(2012).Mutations in the Arabidopsis homolog of LST8/GβL,a partner of the Target of Rapamycin kinase,impair plant growth,flowering,and metabolic adaptation to long days.Plant Cell,24(2):463-481
Peng Y,Chen L,Li S,et al(2018).BRI1 and BAK1 interact with G proteins and regulate sugar-responsive growth and development in Arabidopsis.Nat Commun,9(1):1522
Riou-Khamlichi C,Menges M,Healy JM,et al(2000).Sugar control of the plant cell cycle:differential regulation of Arabidopsis D-type cyclin gene expression.Mol Cell Biol,20(13):4513-4521
Robaglia C,Thomas M,Meyer C(2012).Sensing nutrient and energy status by SnRK1 and TOR kinases.Curr Opin Plant Biol,15(3):301-307
Roldan M,Gomez-Mena C,Ruiz-Garcia L,et al(1999).Sucrose availability on the aerial part of the plant promotes morphogenesis and flowering of Arabidopsis in the dark.Plant J,20(5):581-590
Schoof H,Lenhard M,Haecker A,et al(2000).The stem cell population of Arabidopsis shoot meristems in maintained by a regulatory loop between the CLAVATA and WUS-CHEL genes.Cell,100(6):635-644
Skylar A,Sung F,Hong F,et al(2011).Metabolic sugar signal promotes Arabidopsis meristematic proliferation via G2.Dev Biol,351(1):82-89
Smeekens S,Ma J,Hanson J,et al(2010).Sugar signals and molecular networks controlling plant growth.Curr Opin Plant Biol,13(3):274-279
Takagi H,Abe A,Yoshida K,et al(2013).QTL-seq:rapid mapping of quantitative trait loci in rice by whole genome resequencing of DNA from two bulked populations.Plant J,74(1):174-183
Timson D(2007).GHMP kinases--structures,mechanisms and potential for therapeutically relevant inhibition.Curr Enzym Inhib,3(1):77-94
Tsai AY,Gazzarrini S(2014).Trehalose-6-phosphate and SnRK1 kinases in plant development and signaling:the emerging picture.Front Plant Sci,5:1-11
UniProt Consortium(2015).UniProt:a hub for protein information.Nucleic Acids Res,43:204-212
Wang G,Ellendorff U,Kemp B,et al(2008).A genome-wide functional investigation into the roles of receptor-like proteins in Arabidopsis.Plant Physiol,147(2):503-517
Wang Y,Li J(2008).Molecular basis of plant architecture.Annu Rev Plant Biol,59:253-279
Wolf S,Ladwig F,Sticht C,et al(2014).A receptor-like protein mediates the response to pectin modification by activating brassinosteroid signaling.Proc Natl Acad Sci USA,111(42):15261-15266
Wu J,Liu Z,Zhang Z,et al(2016).Transcriptional regulation of receptor-like protein genes by environmental stresses and hormones and their overexpression activities in Arabidopsis thaliana.J Exp Bot,67(11):3339-3351
Wu X,Dabi T,Weigel D(2005).Requirement of homeobox gene STIMPY/WOX9 for Arabidopsis meristem growth and maintenance.Curr Biol,15(5):436-440
Xiong F,Zhang R,Meng Z,et al(2017).Brassinosteriod Insensitive 2(BIN2)acts as a downstream effector of the Target of Rapamycin(TOR)signaling pathway to regulate photoautotrophic growth in Arabidopsis.New Phytol,213(1):233-249
Xiong Y,Sheen J(2012).Rapamycin and glucose-target of rapamycin(TOR)protein signaling in plants.J Biol Chem,287(4):2836-2842
Yang T,Bar-Peled L,Gebhart L,et al(2009).Identification of galacturonic acid-1-phosphate kinase,a new member of the GHMP kinase superfamily in plants,and comparison with galactose-1-phosphate kinase.J Biol Chem,284(32):21526-21535
Zhang Y,Liu Z,Wang J,et al(2015).Brassinosteroid is required for sugar promotion of hypocotyl elongation in Arabidopsis in darkness.Planta,242(4):881-893
Zhang Z,Zhu JY,Roh J,et al(2016).TOR signaling promotes accumulation of BZR1 to balance growth with carbon availability in Arabidopsis.Curr Biol,26(14):1854-1860
Zheng L,Shang L,Chen X,et al(2015).TANG1,encoding a Symplekin_C domain-contained protein,influences sugar responses in Arabidopsis.Plant Physiol,168(3):1000-1012