楊淑華

1994年7月－1998年6月，任南開大學助理研究員。

2002－2004年，在美國康奈爾大學植物生物系從事博士后研究。

2005年1月至今，中國農業大學生物學院植物系教授。

研究方向

植物響應低溫脅迫的分子機制。

作為外界主要環境因子，溫度影響植物的生長發育、植被的分佈以及農作物的產量。我們實驗室主要是通過分子生物學、遺傳學和生物化學等手段，研究植物應答低溫脅迫的信號轉導途徑、低溫信號與植物激素信號之間的交互作用，以及低溫信號與生物脅迫信號之間的交互作用等。

1.植物響應零下低溫脅迫的分子調控研究

在植物抗凍性研究方面，我們建立了穩定的凍相關突變體的篩選體系。通過該體系，我們發現蛋白激酶CRPK1（Cold-responsive protein kinase 1）參與負向調控低溫信號途徑關鍵轉錄因子CBF依賴的信號轉導。低溫激活定位於細胞膜上的CRPK1，後者磷酸化細胞質中的14-3-3蛋白，誘導其進入細胞核中，與CBF1/3蛋白互作，介導CBF1/3蛋白的泛素化降解，從而負調控植物對低溫的應答。該研究不僅揭示了低溫信號從細胞膜傳遞到細胞核中的負調控機制，也闡明了CBF蛋白翻譯后修飾的調控機制（Liu et al., 2017,Mol Cell）。而蛋白激酶OST1在植物低溫應答中發揮重要的正向調控作用。OST1激酶活性被低溫迅速誘導，激活狀態的OST1將ICE1磷酸化，抑制ICE1的泛素化降解，從而正調控植物的抗凍性（Ding et al., 2015,Dev Cell）。

在低溫信號和植物激素信號的交叉互作方面，我們發現乙烯信號通路關鍵組分均參與低溫應答。轉錄因子EIN3蛋白直接負調控低溫調節基因CBF和細胞分裂素信號負調控因子A型ARR基因的表達，從而負調控擬南芥的抗凍性。該研究明確了乙烯在植物抗低溫中的作用，同時也揭示乙烯和細胞分裂素信號之間交互作用的分子機制（Shi et al., 2012,Plant Cell）。另一個重要植物激素油菜素內酯信號通路關鍵的蛋白激酶BIN2和轉錄因子BZR1/BES1通過依賴和不依賴於CBF的方式調控下游一系列基因的表達，從而正向調節植物的抗凍能力（Li et al., 2017,Mol Plant）。利用CRISPR技術，構建了cbfs三缺失突變體，明確了CBFs在低溫馴化中的關鍵作用及CBFs的靶基因，並為低溫信號研究提供了理想的遺傳材料 (Jia et al., 2016,New Phytol)。

2.植物響應零上低溫脅迫的分子調控研究

我們的研究發現擬南芥防衛反應中起重要作用的抗病相關基因CHS1、RPP4/CHS2、CHS3和LSD1/CHS4參與不同溫度下植物生長的穩態平衡調控 (Wang et al., 2013,Plant J;Wang et al., 2011,Plant J;Huang et al., 2010,Plant Physiol; Yang et al., 2010,Plant J; Huang et al., 2010,New Phytol)。研究揭示TIR-NB類蛋白CHS1通過與R蛋白SOC3互作，蛋白磷酸酶IBR5和伴侶蛋白HSP90分別與R蛋白CHS3和CHS2互作，調控R蛋白的活性，從而調控R介導的溫度依賴的細胞死亡和防衛反應 (Zhang et al., 2016,New Phytol; Liu et al., 2015,PLoS Genet; Bao et al., 2014,New Phytol)。

研究發現擬南芥磷脂結合蛋白BON1在調控溫度依賴的生長發育中是一個關鍵調控因子，BON1與兩個類受體蛋白激酶BIR1和BAK1互作，可能通過BAK1對BIR1和BON1的磷酸化，作用於同一條信號通路，協同調控溫度依賴的植物生長發育和防衛反應 (Wang et al., 2011,Plant J; Yang et al.,2007,Plant Physiol; Yang et al., 2006,Plant J; Yang and Hua, 2004,Plant Cell)。

論文

1.Liu Z, Jia Y, Ding Y, Shi Y, Zhang X, Li Z, Guo Y, Gong Z,Yang S*(2017) Plasma membrane CRPK1-mediated phosphorylation of 14-3-3 proteins induces their nuclear import to fine-tune CBF signaling during cold response.Mol Cell66: 117-128. (Previewed in the same issue of Mol Cell)

2.Li H#, Ye K#, Shi Y, Zhang X,Yang S*(2017) BZR1 positively regulates freezing tolerance via CBF-dependent and CBF-independent pathways in Arabidopsis.Mol Plant 10: 545-559. (#These authors contributed equally) (Spotlighted in the same issue of Mol Plant)

3.Zhang Z#, Li J#, Pan Y#, Li J#, Zhou L#, Shi H#, Zeng Y#, Guo H, Yang S, Zheng W, Yu J, Sun X, Li G, Ding Y, Ma L, Shen S, Dai Ln, Zhang H,Yang S, Guo Y, Li Z* (2017) Natural variation in CTB4a enhances rice adaptation to cold habitats.Nat Commun8:14788. Doi: 10.1038/ncomms14788.

4.Zhang Y#, Wang Y#, Liu J, Ding Y, Wang S, Zhang X, Liu Y,Yang S*(2017) Temperature-dependent autoimmunity mediated by chs1 requires its neighboring TNL gene SOC3.New Phytol213:1330-1345.

5.Jia Y, Ding Y, Shi Y, Zhang X, Gong Z,Yang S*(2016) The cbfs triple mutants reveal the essential functions of CBFs in cold acclimation and allow the definition of CBF regulons in Arabidopsis.New Phytol212: 345-353. (Commentary in the same issue of New Phytol)

6.Huang X, Zhang X, Gong Z,Yang S, Shi Y* (2016) ABI4 represses the expression of type-A ARRs to inhibit seed germination in Arabidopsis.Plant J89: 354-365.

7.Yu F, Lou L, Tian M, Li Q, Ding Y, Wu Y, Belda-Palazon B, Rodrigue P L,Yang S, Xie Q* (2016) ESCRT-I component VPS23A affects ABA signaling by recognizing ABA receptors for endosomal degradation.Mol Plant9: 1570-1582.

8.Ding Y, Li H, Zhang X, Xie Q, Gong Z,Yang S* (2015) OST1 kinase modulates freezing tolerance by enhancing ICE1 stability in Arabidopsis.Dev Cell 32: 278-289. (Previewed in the same issue of Mol Cell)

9.Liu J#, Yang H#, Bao F, Ao K, Zhang X, Zhang Y,Yang S* (2015) IBR5 modulates temperature-dependent, R protein CHS3-mediated defense responses in Arabidopsis.PLoS Genet11, e1005584. (#These authors contributed equally)

10.Kong L, Cheng J, Zhu Y, Ding Y, Meng J, Chen Z, Xie Q, Guo Y, Li J,Yang S, Gong Z* (2015) Degradation of the ABA co-receptor ABI1 by PUB12/13 U-box E3 ligases.Nat Commun6: 8630. Doi: 10.1038/ncomms9630.

11.Ji H, Wang Y, Cloix C, Li K, Jenkin G, Wang S, Shang Z, Shi Y,Yang S, Li X* (2015) The Arabidopsis RCC1 family protein TCF1 regulates freezing tolerance and cold acclimation through modulating lignin biosynthesis.PLoS Genet11, e1005471.

12.Shi Y#, Ding Y#, Yang S* (2015) Cold signal transduction and its interplay with phytohormones during cold acclimation. Plant Cell Physiol56: 7-15. (#These authors contributed equally)

13.Bao F, Huang X, Zhu C, Zhang X, Li X, Yang S*(2014) Arabidopsis HSP90 protein modulates RPP4-mediated temperature-dependent cell death and defense responses. New Phytol202: 1320-1334.

14.Song S, Huang H, Gao H, Wang J, Wu D, Liu X, Yang S, Zhai Q, Li C, Qi T, Xie D* (2014) Interaction between MYC2 and ETHYLENE INSENSITIVE3 modulates antagonism between jasmonate and ethylene signaling in Arabidopsis. Plant Cell26: 263-279.

15.Wang Y, Zhang Y, Wang Z, Zhang X,Yang S*(2013) A missense mutation in CHS1, a TIR-NB protein, induces chilling sensitivity in Arabidopsis. Plant J 75: 553-565.

16.Shi Y, Wang Z, Meng P, Tian S, Zhang X, Yang S* (2013) The glutamate carboxypeptidase AMP1 mediates ABA and abiotic stress responses in Arabidopsis. New Phytol 199: 135-150.

17.Guo L, Yang H, Zhang X, Yang S*(2013) Lipid transfer protein 3 as a target of MYB96 mediates freezing and drought stress in Arabidopsis. J Exp Bot64: 1755-1767.

18.Li J#, Mu J#, Bai J#, Fu F, Zou T, An F, Zhang J, Jing H, Wang Q, Yang S, Zuo J* (2013) PARAQUAT RESISTANT 1, a Golgi-localized putative transporter protein, is involved in intracellular transport of paraquat. Plant Physiol 162: 470-483. (#These authors contributed equally)

19.Shi Y#, Tian S#, Hou L, Huang X, Zhang X, Guo H, Yang S*(2012) Ethylene signaling negatively regulates freezing tolerance by repressing expression of CBF and type-A ARR genes in Arabidopsis. Plant Cell24: 2578-2595. (#These authors contributed equally)

20.Wang Z#, Meng P#, Zhang X, Ren D,Yang S*(2011) BON1 interacts with the protein kinases BIR1 and BAK1 in modulation of temperature-dependent plant growth and cell death in Arabidopsis.Plant J 67: 1081-1093. (#These authors contributed equally)

21.Huang X, Li J, Bao F, Zhang X,Yang S*(2010) A gain-of-function mutation in the Arabidopsis disease resistance gene RPP4 confers sensitivity to low temperature.Plant Physiol154: 796-809.

22.Yang H, Shi Y, Liu J, Guo L, Zhang X,Yang S*(2010) A mutant CHS3 protein with TIR-NB-LRR-LIM domains modulates growth, cell death and freezing tolerance in a temperature-dependent manner in Arabidopsis.Plant J63: 283-296.

23.Huang X, Li Y, Zhang X, Zuo J,Yang S*(2010) The Arabidopsis LSD1 gene plays an important role in the regulation of low temperature-dependent cell death.New Phytol 187: 301-312 (Cover story).

24.Liu D, Gong Q, Ma Y, Li P, Li J,Yang S, Yuan L, Yu Y, Pan D, Xu F, Wang NN* (2010) cpSecA, a thylakoid protein translocase subunit, is essential for photosynthetic development in Arabidopsis.J Exp Bot 61: 1655-1669.

25.Huang X, Zhang X,Yang S*(2009) A novel chloroplast-localized protein EMB1303 is required for chloroplast development in Arabidopsis.Cell Res19: 1205-1216.

26.Gou M, Su N, Zheng J, Huai J, Wu G, Zhao J, He J, Tang D,Yang S, Wang G* (2009) An F-box gene, CPR30, functions as a negative regulator of the defense response in Arabidopsis.Plant J 60: 757-770.

27.Li Y#,Yang S#, Yang H, Hua J* (2007) The TIR-NB-LRR gene SNC1 is regulated at the transcript level by multiple factors.Mol Plant Microbe Interact20: 1449-1456. (#These authors contributed equally)

28.Yang H,Yang S, Li Y, Hua J* (2007) The Arabidopsis BAP1 and BAP2 genes are general inhibitors of programmed cell death.Plant Physiol 145: 135-146.

29.Chai MF, Wei PC, Chen QJ, An R, Chen YM, Chen J,Yang S*, Wang XC* (2006) NADK3, a novel cytoplasmic source of NADPH, is required under conditions of oxidative stress and modulates abscisic acid responses in Arabidopsis.Plant J47: 665-674.

30.Yang S, Yang H, Grisafi P, Sanchatjate S, Fink GR, Sun Q, Hua J* (2006) The BON/CPN gene family represses cell death and promotes cell growth in Arabidopsis.Plant J 45: 166-179.

31.YangS, and Hua J* (2004) A haplotype-specific resistance gene regulated by BONZAI1 mediates temperature-dependent growth control in Arabidopsis.Plant Cell16: 1060-1071.

32.Yang S*, Yu H, Xu Y, Goh CJ* (2003) Investigation of cytokinin-deficient phenotypes in Arabidopsis by ectopic expression of orchid DSCKX1.FEBS Lett 555: 291-296.

33.Yang S, Yu H, Goh CJ* (2003) Functional characterisation of a cytokinin oxidase gene DSCKX1 in Dendrobium orchid. Plant Mol Biol 51: 237-248.

34.Yang S, Yu H, Goh CJ* (2002) Isolation and characterization of the orchid cytokinin oxidase DSCKX1 promoter.J Exp Bot 53: 1899-1907.

35.Yu H,Yang S, Goh CJ* (2002) Spatial and temporal expression of the orchid floral homeotic gene DOMADS1 is mediated by its upstream regulatory elements.Plant Mol Biol49: 225-237.

36.Yu H,Yang S, Goh CJ* (2001) Agrobacterium-mediated transformation of a Dendrobium orchid with the class 1 knox gene DOH1.Plant Cell Rep 20: 301-305.

37.Yu H,Yang S, Goh CJ* (2000) DOH1, a class 1 knox Gene, is required for maintenance of the basic plant architecture and floral transition in orchid.Plant Cell 12: 2143-2160.

38.丁楊林，施怡婷，楊淑華(2015)植物響應低溫脅迫的分子機制研究。生命科學.27:398-405.

39.鄧岩，王興春，楊淑華，左建儒(2006) 細胞分裂素：代謝、信號轉導、交叉反應與農藝性狀改良. 植物學通報.23:478-498.

專著

Shi Y,Yang S*(2014) ABA regulation of the cold stress response in plants. In Zhang D (ed)Abscisic Acid: Metabolism, Transport and Signaling.Springer, pp337-363.

Yu H, Dhavale T,Yang S(2006) Molecular mechanisms of hormone functions in flowering In Teixeira da Silva JA (ed) Floriculture, Ornamental and Plant Biotechnology: Advances and Topical Issues, Ed 1 Vol 1 Global Science Books, London, UK, pp 25-32.

任勃，王興春，馮健，楊淑華，左建儒(2012)細胞分裂素。植物激素作用的分子機理(許智宏，薛紅衛主編)上海科學技術出版社.

武維華，張蜀秋，袁明，張軍，韓玉珍，李穎章，葉德，楊淑華，王學臣(2008)植物生理學(第二版)科學出版社.

近年主持國家自然科學基金重大研究計劃、重點項目、面上項目、國家重點基礎研究規劃項目（“973”）子課題等科研項目。

任New Phytologist Editor、Plant & Cell Physiology Advisory Editor、《植物學報》副主編。