Education and Employment

2025-Present: Professor, Shanghai Jiao Tong University

2022-2025: Professor, Shanghai Normal University

1995-2021: Postdoctoral Research Associate, Texas A&M University

1993-1994: Research associate, Shanghai Institute of Plant Physiology (SIPP), CAS, China

1987-1993: Ph. D, SIPP, CAS, China.

1983-1987: Bachelor degree, Nanjiang University, China

Research Interests:

1. Anther Development and Pollen Formation: Identification of genes essential for anther development through male sterile mutants; functional analysis of these genes for the development of each cell layers in the anther (microspore mother cells, tapetal cells, middle layer cells, and endothecium cells), and their relationships with pollen formation at the cellular, genetic, biochemical, and molecular levels. Representative work includes the analysis of the genetic pathway of tapetal cells and the cascade regulatory mechanism for pollen formation. For details, please refer to the review (Yao et al., Seed Biology 1:5, 2022).  

2. Mechanisms of Photo-Thermo Sensitive Male Sterility (P/TGMS) and Two-Line Hybrid Seed Production in Crops: Two-line hybrid rice relies on P/TGMS lines. The shortage of reliable rice P/TGMS lines is currently a bottleneck in hybrid rice breeding. Our research group has elucidated the P/TGMS mechanisms that slow development under low temperature or short light conditions, and the effective scavenging of reactive oxygen species (ROS) are two general factors for P/TGMS fertility restoration. For details, please refer to the review (Wang et al., Molecular Plant 18:26-41, 2025). On this basis, novel P/TGMS lines of rice and maize have been created, among which ostms15, ostms16, ostms18, and ostms19 have been published. ostms15 has been applied in approximately thirty breeding units in China.

3. Mechanisms of Cytoplasmic Male Sterility (CMS) and Fertility Regulation through Nucleo-Cytoplasmic Interaction: CMS plays a crucial role in the three-line hybrid seed production of crops and horticultural plants. We will investigate the mechanisms of CMS at the molecular and cellular levels, and create genetic materials for three-line breeding of crops and horticultural plant.

Publications:

1. Wang N, Li X, Zhu J, Yang ZN*. 2025. Molecular and Cellular Mechanisms of Photoperiod and Thermo-sensitive Genic Male Sterility in Plants. Molecular Plant 18(1):26-41.
2.  
3. Yao XZ, Hu Wen, and Yang ZN*, 2022. The contributions of sporophytic tapetum to pollen formation. Seed Biology 1:5.
4.  
5. Ma C, Wang SH, Yang YM, Fu J, Guo MD, Chen X, Mao YC, Huang XH, Zhu J*, Yang ZN*. 2025. ATP-Citrate Lyase ACLB-1 is critical for sporopollenin synthesis and energy supply for microspore development. Plant Physiology. 197(2)
6.  
7. Zhou L, Mao YC, Yang YM, Wang JJ, Zhong X, Han Y, Zhang YF, Shi QS, Huang XH, Blake C. Meyers, Zhu J*, Yang ZN*. 2024. Temperature and light reverse the fertility of rice P/TGMS line ostms19 via ROS homeostasis. Plant Biotechnology Journal. 22: 2020-2032.
8.  
9. Chen X†, Huang DD†, Xue JS†, Bu JH, Zhang MQ, Wang N, Huang JR, Chen WS* and Yang ZN*. 2024. Polymeric phenylpropanoid derivatives crosslinked by hydroxyl fatty acids form the core structure of rape sporopollenin. Nature Plants 10: 1790-1800 
10.  
11. Zhang C, Xiong AT, Ren MY, Zhao YY, Huang MJ, Huang LC, Zhang Z, Wang Y, Zheng QQ, Fan J, Guan JJ and Yang ZN*. 2024. An epigenetically mediated double negative cascade from EFD to HB21 regulates anther development. Nature Communications 14, 7796.
12.  
13. Zhang Z, Guo YY, Wang YC, Zhou L, Fan J, Mao YC, Yang YM, Zhang YF, Huang XH, Zhu J, Zhang C*, Yang ZN*. 2024. A point mutation in a meiotic crossover formation gene HEI10/TFS2 leads to thermosensitive genic female sterility in rice. Plant Journal 118: 506-518.
14.  
15. Xue JS, Feng YF, Zhang MQ, Xu QL, Xu YM, Shi JQ, Liu LF, Wu XF, Wang S, and Yang ZN*. 2024. The regulatory mechanism of rapid lignification for timely anther dehiscence. JIPB 66(8):1788-1800.
16.  
17. Han Y, Jiang SZ, Zhong X, Chen X, Ma CK, Yang YM, Mao YC, Zhou SD, Zhou L, Zhang YF, Huang XH, Zhang H, Li LG, Zhu J*, Yang ZN*. 2023. Low temperature compensates the defective tapetum initiation to restore the fertility of a novel TGMS line ostms15. Plant Biotechnology Journal 21:1659-1670
18.  
19. Xue JS, Qiu S, Jia XL, Shen SY, Shen CW, Wang S, Xu P, Tong Q, Lou YX, Yang NY, Cao JG, Hu JF, Shen H, Zhu RL, Murray JD, Chen WS*, Yang ZN*. 2023. Stepwise changes in flavonoids in spores/pollen contributed to terrestrial adaptation of plants. Plant Physiology 193(1):627-642
20.  
21. Zhang YF†, Li YL†, Zhong X, Wang JJ, Zhou L, Han Y, Li DD, Wang N, Huang XH, Zhu J*, Yang ZN*. 2022. Mutation of the GMC oxidoreductase leads to thermo-sensitive genic male sterility in rice and Arabidopsis. Plant Biotechnology Journal 20:2023-2035.
22.  
23. Zhang C†, Ren MY†, Han WJ†, Zhang YF, Huang MJ, Wang Y, Zhang Z, Wu SY, Huang J, Yang ZN*. 2022. Slow development allows redundant genes to restore the fertility of rpg1, a TGMS line in Arabidopsis. Plant Journal 109: 1375-1385
24.  
25. Shi QS†, Lou Y†, Shen SY, Wang KQ, Wang SH, Zhou L, Wang JJ, Liu XL, Xiong SX, Han Y, Zhou HS, Huang XH, Wang S, Zhu J*, Yang ZN*. 2021. A cellular mechanism underlying the restoration of thermo/photoperiod-sensitive genic male sterility. Molecular Plant 14:2104-2114.
26.  
27. Zhang C†, Xu T†, Ren MY†, Zhu J, Shi QS, Zhang YF, Qi YW, Huang MJ, Song L, Xu P*, Yang ZN*. 2020. Slow development restores the fertility of photoperiod-sensitive male sterile plant lines. Plant Physiology 184(2): 923-932
28.  
29. Lu JY†, Xiong SX†, Yin WZ†, Teng XD, Lou Y, Zhu J, Zhang C, Gu JN, Wilson ZA*, Yang ZN*. 2020. MS1, a direct target of MS188, regulates the expression of key sporophytic pollen coat protein genes in Arabidopsis thaliana. Journal Experimental Botany 71(16): 4877-4889.  
30.  
31. Xue JS†, Zhang BC†, Zhan HD†, Lv YL, Jia XL, Wang TH, Yang NY, Lou YX, Zhang ZB, Hu WJ, Gui JS, Cao JG, Xu P, Zhou YH, Hu JF, Li LG*, Yang ZN*. 2020. Phenylpropanoid derivatives are essential components of sporopollenin in vascular plants. Molecular Plant 13: 1644-1653
32.  
33. Zhu J†, Lou Y†, Shi QS†, Zhang S†, Zhou WT, Yang J, Zhang C, Yao XZ, Xu T, Liu JL, Wang S, Huang XH, Yang ZN*. 2020. Slowing development restores the fertility of thermo-sensitive male-sterile plant lines. Nature Plants 6: 360-367
34.  
35. Xiong HB, Huang C, Wang J, Rochaix JD, Lin FM, Zhang JX, Ye LS, Shi XH, Yu QB*, and Yang ZN*. 2020. mTERF8, a Member of the Mitochondrial Transcription Termination Factor Family, is Involved in the Transcription Termination of Chloroplast Gene psbJ. Plant Physiology. 182: 408-423.
36.  
37. Xu XF†, Wang B†, Feng YF†, Xue JS, Qian XX, Liu SQ, Zhou J, Yu YH, Yang NY, Xu P, Yang ZN*. 2019. AUXIN RESPONSE FACTOR17 directly regulates MYB108 for anther dehiscence. Plant Physiology 181: 645-655.
38.  
39. Zhang C†, Teng XD†, Zhao YY, Lu JY, Wang YC, Guo HW*, Yang ZN*. 2018. Ethylene signaling control is critical for female gametophyte and embryo development. Plant Journal 96(1): 176-187.
40.  
41. Wang K, Guo ZL, Zhang C,Zhang ZY, Lou Y, Xiong SX, Yao XZ, Fan JJ, Zhu J* and Yang ZN*. 2018. The MYB transcription factor MS188 directly regulates the expressions of sporopollenin synthesis genes for pollen wall formation. Plant Physiology 178: 283-294.
42.  
43. Yao XZ, Tian L, Yang J, Zhao YN, Zhu YX, Dai XH, Zhao YD* & Yang ZN*. 2018. Auxin production in diploid microsporocytes is necessary and sufficient for early stages of pollen development. PLoS Genetics 14 (5): e1007397.
44.  
45. Zhan HD, Xiong HB, Wang S and Yang ZN*. 2018. Anther endothecium provides pollen coat very-long-chain fatty acids (VLCFAs) for pollen hydration in Arabidopsis. Molecular Plant 11(8):1101-1104
46.  
47. Lou Y, Zhou HS, Han Y, Zeng QY, Zhu J*, and Yang ZN*. 2018. Positive regulation of AMS by TDF1 and the formation of a TDF1–AMS complex are required for anther development in Arabidopsis thaliana. New Phytologist 217: 378-391
48.  
49. Huang C†, Yu QB†, Li ZR, Ye LS, Xu L*, and Yang ZN*. 2017. Porphobilinogen deaminase HEMC interacts with the PPR-protein AtECB2 for chloroplast RNA editing. Plant Journal 92(4): 546-556
50.  
51. Yang ZN. 2016. Regulation of sporopollenin synthesis for pollen wall formation in plant. SCIENCE CHINA-LIFE SCIENCES 59(12): 1335-1337.
52.  
53. Xu T†, Zhang C†, Zhou Q and Yang ZN*. 2016. Pollen wall pattern in Arabidopsis. Science Bulletin 61(11): 832-837. (2013CB945100)
54.  
55. Xu XF, Lou Y, Wang B, Lu JY, Li DD, Li LG, Zhu J*, Yang ZN*. 2015. The Magnesium Transporter 5 Plays an Important Role in Mg Transportation from Tapetum to Microspores in Arabidopsis. Plant Journal 84(5):925-936
56.  
57. Cai CF†, Zhu J†, Lou Y, Guo ZL, Xiong SX, Wang K and Yang ZN*. 2015. Thefunctionalanalysisof OsTDF1revealsaconservedgeneticpathwayfor tapetal development between rice and Arabidopsis. Science Bulletin 60(12): 1073-1082
58.  
59. Jia QS, Zhu J, Xu XF, Lou Y, Zhang ZL, Zhang ZP and Yang ZN*. 2015. Arabidopsis AT-hook protein TEK positively regulates the expression of arabinogalactan proteins (AGPs) for nexine formation. Molecular Plant 8:251-260.
60.  
61. Gu JN†, Zhu J†, Yu Y, Teng XT, Lou Y, Xu XF, Liu JL, Yang ZN*. 2014. DYT1 Directly Regulates the Expression of TDF1 for Tapetum Development and Pollen Wall Formation in Arabidopsis. Plant Journal 80: 1005-1013.
62.  
63. Lou Y†, Xu XF†, Zhu J†, Gu JN, Blackmore S, Yang ZN*. 2014. The Tapetal AHL family protein TEK determines nexine formation in the pollen wall. Nature Communications 5:3855
64.  
65. Yu QB, Kong MM, Zhu J, Jia QS, Jiang Y, Mi HL, Chong K, Yang ZN*. 2014. AtECB1/MRL7, a Thioredoxin-like Fold Protein with Disulfide Reductase Activity, Regulates Chloroplast Gene Expression and Chloroplast Biogenesis in Arabidopsis thaliana. Molecular Plant 7(1):206-217
66.  
67. Yang J, Tian L, Huang XY, Sun MX, Zhu J, Guan YF, and Yang ZN*. 2013. Auxin Response Factor17 is essential for auxin signaling in primexine formation and pollen development. Plant Physiology 162:720-731.
68.  
69. Huang XY, Niu J, Sun MX, Zhu J, Gao JF, Yang J, Zhou Q, and Yang ZN*.2013. CDKG1 is Associated with Splicesome to Regulate CalS5 Splicing and Pollen Wall Formation in Arabidopsis thaliana. Plant Cell 25(2) :637-648
70.  
71. Zhang C, Song Y, Cheng ZH, Wang YX, Zhu J, Ma H, Xu L*, Yang ZN*. 2012. The Arabidopsis thaliana DSB FORMATION (AtDFO) Gene is Required for Meiotic Double Strand Break Formation. Plant Journal. 72(2):271-281
72.  
73. Li XC†, Zhu J†, Yang J, Zhang GR, Xing WF, Zhang S, Yang ZN*. 2012. Glycerol-3-phosphate acyltransferase 6 (GPAT6) is important for tapetum development in Arabidopsis and plays multiple roles in plant fertility. Molecular Plant.5(1): 131-142
74.  
75. Chang HS†, Zhang C†, Chang YH, Zhu J, Xu XF, Shi ZH, Zhang XL, Xu L, Huang H, Zhang S*, and Yang ZN*，2012. NO PRIMEXINE AND PLASMA MEMBRANE UNDULATION (NPU), Is Essential for Primexine Deposition and Plasma Membrane Undulation during Microsporogenesis in Arabidopsis thaliana. Plant Physiology 158(1): 264–272.
76.  
77. Gao ZP, Yu QB, Zhao TT, Ma Q, Chen GX*, and Yang ZN*. 2011. A functional component of transcriptionally active chromosome complex, Arabidopsis pTAC14, interacts with pTAC12 and regulates the plastid gene expression. Plant Physiology 157(4): 1733-1745
78.  
79. Cui YL, Jia QS, Yin QQ, Lin GN, Kong MM, Yang ZN*. 2011. The GDC1 Gene Encodes a Novel Ankyrin Domain Containing Protein That is Essential for Grana Formation in Arabidopsis thaliana. Plant Physiology 155(1):130-141.
80.  
81. Zhu J, Lou Y, Xu XF, and Yang ZN*. 2011. A Genetic Pathway for Tapetum Development and Function in Arabidopsis. Journal of Integrative Plant Biology. 53(11): 892-900.
82.  
83. Yu QB†, Jiang Y†, Chong K*, and Yang ZN*. 2009. AtECB2, a Pentatricopeptide Repeat Protein, is Involved in Chloroplast Transcript accD RNA Editing and Essential for Early Chloroplast Biogenesis in Arabidopsis thaliana. Plant Journal. 59(6):1011-1023.
84.  
85. Jiang H†, Wang FF†, Wu YT†, Zhou X, Huang XY, Zhu J, Gao JF, Dong RB，Cao KM*, Yang ZN*.2009. MULTIPOLAR SPINDLE 1 (MPS1), a novel coiled-coil protein of Arabidopsis thaliana, is required for meiotic spindle organization. Plant Journal. 59(6):1001-1010.
86.  
87. Zhu J†, Chen H†, Li H, Gao JF, Jiang H, Wang C, Guan YF, and Yang ZN*. 2008. Defective of Tapetum Development and Function1 (TDF1) Is Essential for Anther Development and Tapetum Function for Microspore Maturation in Arabidopsis. Plant Journal 55(2): 266-277
88.  
89. Guan YF, Huang XY, Zhu J, Gao JF, Zhang HX and Yang ZN*. 2008. RUPTURED POLLEN GRAIN 1 (RPG1), a Member of MtN3/Saliva Gene Family, is Crucial for Exine Pattern Formation and Cell Integrity of Microspores in Arabidopsis thaliana. Plant Physiology 147(2): 852-862
90.  
91. Yu QB†, Li G†, Wang G†, Sun JC†, Wang PC, Wang C, Mi HL, Ma WM, Cui J, Cui YL,Chong K, Li YX, Li YH, Zhao ZM, Shi TL*, and Yang ZN*. 2008. Construction of a Chloroplast Protein Interaction Network and Functional Mining of Photosynthetic Proteins in Arabidopsis thaliana. Cell Research. 18(10):1007-1019.
92.  
93. Zhang ZB†, Zhu J†, Gao JF†, Wang C, Li H, Zhang HQ, Zhang S, Wang DM, Wang QX, Huang H, Xia HJ, Yang ZN*. 2007. Transcription Factor AtMYB103 is Required for Anther Development by Regulating Microspore Release from Tetrads and Exine Formation in Arabidopsis. Plant Journal 52(3): 528-538。
94.  
95. Shen YJ†, Jiang H†, Jin JP†, Zhang ZB, Xi B, Wang G, Wang C, Qian L, Li X, Yu QB, Liu HJ, Chen DH, Gao JH, Huang H, Shi TL* & ZN Yang*. 2004. Development of genome-wide DNA polymorphism database for map-based cloning of rice genes. Plant Physiology 135：1198-1205