UV-B对设施油桃叶片叶绿素生物合成的影响

doi:10.16420/j.issn.0513-353x.2023-0482

摘要/Abstract

摘要：

为明晰桃叶片叶绿素生物合成过程中对紫外线B（Ultraviolet B，UV-B）辐照响应的关键步骤，以设施栽培的‘中油5号’油桃（Prunus persica var. nectariana）7年生树为试材，自花后7 d开始，每天8：30—9：30补充适宜剂量（由前期试验筛出，约1.44 kJ · m^-2· d^-1）的UV-B直至果实采收，以正常生长、未补充UV-B辐照的植株为对照，对生育期内叶片叶绿素生物合成过程中前体物质、中间产物和叶绿素含量以及关键酶活性的动态变化进行了探究。结果表明，在叶绿素合成第一阶段中，UV-B辐照通过提升前体物质δ-氨基酮戊酸（ALA）合成关键酶——谷氨酸酯-1-半醛 2,1-氨基变位酶（GSA-AM）的活性，增加了ALA的积累量，为生物合成的开始奠定物质基础。在第二阶段中，UV-B提高了胆色素原脱氨酶（PBGD）的活性，促进了羟甲基胆色素原的合成，为后续产物尿卟啉原Ⅲ（Urogen Ⅲ）的合成提供底物，使得整个生育期内叶片中粪卟啉原Ⅲ（Coprogen Ⅲ）和原卟啉Ⅸ（Proto Ⅸ）的含量显著提升。UV-B辐照后第三阶段的Mg-螯合酶（MgCH）和Mg-原卟啉Ⅸ单甲基酯环化酶（MgPEC）等关键酶的活性显著升高，各中间产物Mg-原卟啉Ⅸ（MgP Ⅸ）、Mg-原卟啉Ⅸ甲酯（MgPME）、联乙烯原叶绿素酸酯、原叶绿素酸酯（Pchlide）、叶绿素酸酯a（Chlide a）、叶绿素酸酯b（Chlide b）含量均得到提升，最后产物叶绿素的含量也显著高于对照。可见UV-B辐照影响叶绿素生物合成的主要关键步骤为前体物质ALA的合成、羟甲基胆色素原的合成、Urogen Ⅲ向Coprogen Ⅲ的转化、Mg²⁺插入Proto Ⅸ及其后等4个步骤，且对叶绿素生物合成的影响具有长期积累性。

关键词: 油桃, 设施栽培, 紫外线B, 叶绿素, 生物合成

Abstract:

In order to clarify the key steps in the response of chlorophyll（Chl）biosynthesis to ultraviolet B（UV-B）irradiation in greenhouse nectarine leaves，the 7-year-old nectarine‘Zhongyou 5’was used as the test material，which was cultivated in greenhouse from 7 days after flowering. The appropriate dose of UV-B（about 1.44 kJ · m^-2· d^-1 screened by previous experiments）was supplemented at 8：30—9：30 every morning until the fruit was harvested. The dynamic changes of precursor substances，intermediate product content，Chl content and key enzyme activity in the process of Chl biosynthesis in leaves during the growth and development period were investigated with normal growth and unsupplemented UV-B irradiated plants as the control. The results showed that in the first stage of Chl synthesis，UV-B irradiation increased the accumulation of δ-aminolevulinic acid（ALA）by increasing the activity of glutamate-1-semialdehyde-2,1-aminomutase（GSA-AM），a key enzyme in the synthesis of precursor ALA，which laid a good material foundation for the beginning of biosynthesis. In the second stage，UV-B increased the activity of bile pigment deaminase（PBGD），promoted the synthesis of hydroxymethyl bile pigment，and provided sufficient substrates for the synthesis of subsequent products uroporphyrinogen Ⅲ（Urogen Ⅲ），so that the content of Coprogen Ⅲ and Proto Ⅸ in leaves increased significantly during the whole growth period. After UV-B irradiation，the activities of key enzymes such as Mg-chelating enzyme（MgCH）and Mg-protoporphyrin Ⅸ monomethyl ester cyclase（MgPEC）in the third stage were significantly increased，and the content of intermediate products Mg-protoporphyrin Ⅸ（MgPⅨ），Mg-protoporphyrin Ⅸ methyl ester（MgPME），bivinyl protochlorophyllate，protochlorophyllate（Pchlide），chlorophyllate a（Chlide a）and chlorophyllate b（Chlide b）were all increased，the chlorophyll content of the product was also significantly higher than that of the control. It can be seen that the main key steps of UV-B irradiation affecting Chl biosynthesis are the synthesis of precursor ALA，the synthesis of hydroxymethylcholanogen，the conversion of Urogen Ⅲ to Coprogen Ⅲ，the insertion of Mg^{2 +} into Proto Ⅲ and the following four steps，and the effect of UV-B on Chl biosynthesis has a long-term accumulation.

Key words: nectarine, facility cultivation, UV-B, chlorophyll, biosynthesis

田歌, 刘建廷, 高传彩, 赵雪惠, 樊永信, 李森, 张寒啸, 陈修德, 李玲, 李冬梅. UV-B对设施油桃叶片叶绿素生物合成的影响[J]. 园艺学报, 2024, 51(6): 1332-1344.

TIAN Ge, LIU Jianting, GAO Chuancai, ZHAO Xuehui, FAN Yongxin, LI Sen, ZHANG Hanxiao, CHEN Xiude, LI Ling, LI Dongmei. Effects of UV-B on Chlorophyll Biosynthesis in Greenhouse Nectarine Leaves[J]. Acta Horticulturae Sinica, 2024, 51(6): 1332-1344.

图/表 7

图1 UV-B处理对油桃叶片中“谷氨酸→ALA”合成的影响 *P < 0.05；**P < 0.01. 下同。The same below.

Fig. 1 Effects of UV-B treatment on the synthesis of glutamate to ALA in nectarine leaves

图2 UV-B处理对油桃叶片中“ALA→原卟啉Ⅸ”合成的影响

Fig. 2 Effects of UV-B treatment on the synthesis of ALA to Proto Ⅸ in nectarine leaves

图3 UV-B处理对油桃叶片中“原卟啉Ⅸ→叶绿素”中间产物的影响

Fig. 3 Effects of UV-B treatment on the synthesis of Proto Ⅸ to Chl in nectarine leaves

图4 UV-B处理对油桃叶片叶绿素含量的影响

Fig. 4 Effects of UV-B treatment on Chl content in nectarine leaves

图5 UV-B处理对油桃叶片GluTR和GSA-AM活性的影响

Fig. 5 Effects of UV-B treatment on the activities of GluTR and GSA-AM in nectarine leaves

图6 UV-B处理对油桃叶片PBGD和MgCH活性的影响

Fig. 6 Effects of UV-B treatment on the activity of PBGD and MgCH in nectarine leaves

图7 UV-B处理对油桃叶片MgPEC、DVR、POR和CAO活性的影响

Fig. 7 Effects of UV-B treatment on the activities of MgPEC，DVR，POR and CAO in nectarine leaves

参考文献 45

[1]	Beale S I. 2005. Green genes gleaned. Trends in Plant Science, 10 (7):309-312. pmid: 15951223
[2]	Chen Hui-ze, Han Rong. 2015. Plants respond to UV-B radiation:a review. Chinese Bulletin of Botany, 50 (6):790-801. (in Chinese)
	陈慧泽, 韩榕. 2015. 植物响应UV-B辐射的研究进展. 植物学报, 50 (6):790-801. doi: 10.11983/CBB14176
[3]	Cui Xi-tian, Yuan Feng-hui, Wang An-zhi, Guan De-xin, Wu Jia-bing, Jin Chang-jie. 2017. Leaf age-related changes in photosynthesis of Quercus mongolica leaves in relation to leaf functional traits. Chinese Journal of Ecology, 36 (11):3160-3167. (in Chinese)
	崔西甜, 袁凤辉, 王安志, 关德新, 吴家兵, 金昌杰. 2017. 蒙古栎叶片光合作用随叶龄的变化及其与叶片功能性状的关系. 生态学杂志, 36 (11):3160-3167.
[4]	Czarnecki O, Grimm B. 2012. Post-translational control of tetrapyrrole biosynthesis in plants,algae,and cyanobacteria. Journal of Experimental Botany, 63 (4):1675-1687. doi: 10.1093/jxb/err437 pmid: 22231500
[5]	Fan Yong-xin, Zhang Han-xiao, Gao Chuan-cai, Li Hui-ling, Fu Xi-ling, Xiao Wei, Li Ling, Chen Xiu-de, Li Dong-mei. 2023. Effects of UV-B irradiation on growth and IAA metabolism of greenhouse nectarine. Plant Physiology Journal, 59 (2):345-352. (in Chinese)
	樊永信, 张寒啸, 高传彩, 李慧玲, 付喜玲, 肖伟, 李玲, 陈修德, 李冬梅. 2023. UV-B辐照对设施油桃生长及IAA代谢的影响. 植物生理学报, 59 (2):345-352.
[6]	Guo Jian-jun, Gui Meng-yuan, Zhang Pei-pei, Wang Jing, Wang Han-bo, Li Jing-yuan. 2017. Sensitivity of Sedum aizoon to the enhanced ultraviolet B radiation. Journal of Henan Agricultural University, 51 (1):71-75,81. (in Chinese)
	郭建军, 贵梦园, 张佩佩, 王静, 王瀚博, 李景原. 2017. 景天三七对紫外线B辐射敏感性研究. 河南农业大学学报, 51 (1):71-75,81.
[7]	Guo Yu-cai. 2009. Effects of film and UV-B on morphology and leaf physiology of peach tree in greenhouse[M. D. Dissertation]. Tai’an: Shandong Agricultural University. (in Chinese)
	郭玉才. 2009. 不同棚膜及人工补充紫外线B对设施桃树形态建造和叶片生理的影响[硕士论文]. 泰安: 山东农业大学.
[8]	Hailemichaeln G, Catalina A, Gonzalez M R, Martin P. 2016. Relationships between water status,leaf chlorophyll content and photosynthetic performance in Tempranillo vineyards. South African Journal of Enology and Viticulture, 37 (2):149-156.
[9]	Hu J F, Fang H C, Wang J, Yue X X, Su M Y, Mao Z L, Zou Q, Jiang H Y, Guo Z W, Yu L, Feng T, Lu L, Peng Z G, Zhang Z Y, Wang N, Chen X S. 2020. Ultraviolet B-induced MdWRKY72 expression promotes anthocyanin synthesis in apple. Plant Science, 292:110377.
[10]	Inostroza-Blancheteau C, Acevedo P, Loyola R, Arce-Johnson P, Alberdi M, Reyes-Díaz M. 2016. Short-term UV-B radiation affects photosynthetic performance and antioxidant gene expression in highbush blueberry leaves. Plant Physiology and Biochemistry, 107:301-309. doi: S0981-9428(16)30240-6 pmid: 27343876
[11]	Li Chen, Liu Jian-ting, Fan Yong-xin, Zhao Xue-hui, Xiao Wei, Chen Xiu-de, Fu Xi-ling, Li Ling, Li Dong-mei. 2022. Effects of UV-B on Photosynthetic function and chloroplast ultrastructure of peach leaves grown in greenhouse. Chinese Bulletin of Botany, 57 (4):434-443. (in Chinese)
	李晨, 刘建廷, 樊永信, 赵雪惠, 肖伟, 陈修德, 付喜玲, 李玲, 李冬梅. 2022. UV-B对设施桃叶片光合功能及叶绿体超微结构的影响. 植物学报, 57 (4):434-443. doi: 10.11983/CBB21136
[12]	Li Chen, Zhao Xue-hui, Wang Qing-jie, Wang Xu-xu, Xiao Wei, Chen Xiu-de, Fu Xi-ling, Li Ling, Li Dong-mei. 2019. Genome identification of PpGRAS family and expression pattern analysis of responding to UV-B in peach. Scientia Agricultura Sinica, 52 (24):4567-4581. (in Chinese) doi: 10.3864/j.issn.0578-1752.2019.24.011
	李晨, 赵雪惠, 王庆杰, 王旭旭, 肖伟, 陈修德, 付喜玲, 李玲, 李冬梅. 2019. 桃GRAS家族全基因组鉴定与响应UV-B的表达模式分析. 中国农业科学, 52 (24):4567-4581.
[13]	Li Dong-mei, Li Shao-xuan, Xu Gong-xun, Li Chen, Fu Xi-ling, Chen Xiu-de, Zhang Hai-sen, Gao Dong-sheng. 2018. Research advances of plant response to UV-B radiation in greenhouse. Plant Physiology Journal, 54 (1):36-44. (in Chinese)
	李冬梅, 李少旋, 徐功勋, 李晨, 付喜玲, 陈修德, 张海森, 高东升. 2018. 设施作物响应UV-B辐射的研究进展. 植物生理学报, 54 (1):36-44.
[14]	Li Gen, Zhang Cheng, Wang Qiang, Wang Ke, Liu Si-xi, Yang Xun,Wu Ji-kai,Qing Qiu-jing. 2021. Chlorophyll metabolic pathway and its molecular regulation in plants. Science and Technology of Sichuan Agriculture,(4):41-45. (in Chinese)
	李根, 张成, 王强, 王科, 刘思汐, 杨勋, 吴继开, 卿秋静. 2021. 植物叶绿素代谢途径及其分子调控. 四川农业科技,(4):41-45.
[15]	Li Jia-jia, Yu Xu-dong, Cai Ze-ping, Wu Fan-hua, Luo Jia-jia, Zheng Li-ting, Chu Wen-qing. 2019. An overview of chlorophyll biosynthesis in higher plants. Molecular Plant Breeding, 17 (18):6013-6019. (in Chinese)
	李佳佳, 于旭东, 蔡泽坪, 吴繁花, 罗佳佳, 郑李婷, 楚文清. 2019. 高等植物叶绿素生物合成研究进展. 分子植物育种, 17 (18):6013-6019.
[16]	Li Su, Wei Wei,Zhu Yan-lei,Du Juan. 2017. Structural changes of grape leaves and pericarp under ultraviolet light(UVB)stress. Contemporary Horticulture,(6):5-7. (in Chinese)
	李甦, 魏薇, 朱燕蕾, 杜娟. 2017. 紫外线(UVB)胁迫下葡萄叶片和果皮结构变化探究. 现代园艺,(6):5-7.
[17]	Li Tianlai, Qi Mingfang, Meng Sida. 2022. Sixty years of facility horticulture development in China:achievements and prospects. Acta Horticulturae Sinica, 49 (10):2119-2130. doi: 10.16420/j.issn.0513-353x.2022-0700
[18]	李天来, 齐明芳, 孟思达. 2022. 中国设施园艺发展60年成就与展望. 园艺学报, 49 (10):2119-2130. doi: 10.16420/j.issn.0513-353x.2022-0700
[19]	Li T S, Yamane H, Tao R. 2021. Preharvest long-term exposure to UV-B radiation promotes fruit ripening and modifies stage-specific anthocyanin metabolism in highbush blueberry. Horticulture Research, 8 (1):67. doi: 10.1038/s41438-021-00503-4 pmid: 33790261
[20]	Li Yu-hong, Li Gui-rong, Yang Wen-jun, Meng Yan, Wang Jun-zhe, Zhang Cheng-lin. 2022. Construction of Corynebacterium glutamicum for synthesis of 5-aminolevulinic acid. Food and Fermentation Industries, 48 (24):8-15. (in Chinese)
	李宇虹, 李贵荣, 杨文君, 孟燕, 王浚哲, 张成林. 2022. 合成5-氨基乙酰丙酸谷氨酸棒杆菌的代谢工程构建. 食品与发酵工业, 48 (24):8-15. doi: 10.13995/j.cnki.11-1802/ts.031426
[21]	Liu M, Zhao J Q, Pang Y J, Zhang L P, Bian F H, Li L X. 2022. UV-B irradiation and allelopathy by Sargassum thunbergii affects the activities of antioxidant enzymes and their isoenzymes in Corallina pilulifera. Journal of Oceanology and Limnology, 40 (5):1950-1962.
[22]	Liu Min, Li Rong-gui, Fan Hai,Du Gui-cai. 2007. Effects of enhanced UV-B radiation on photosynthetic pigments and some enzymes in tobacco. Acta Botanica Boreali-Occidentalia Sinica, 27 (2):291-296. (in Chinese)
	刘敏, 李荣贵, 范海, 杜桂彩. 2007. UVB辐射对烟草光合色素和几种酶的影响. 西北植物学报, 27 (2):291-296.
[23]	Liu Yi-nuo, Ao Man, Li Bo, Guan Yi-xin. 2020. Effect of ultraviolet-B(UV-B)radiation on plant growth and development and its application value. Soils and Crops, 9 (2):191-202. (in Chinese)
	刘一诺, 敖曼, 李波, 关义新. 2020. UV-B辐射对植物生长发育的影响及其应用价值. 土壤与作物, 9 (2):191-202.
[24]	Llorens L, Badenes-Pérez F R, Julkunen-Tiitto R, Zidorn C, Fereres A, Jansen M A K. 2015. The role of UV-B radiation in plant sexual reproduction. Perspectives in Plant Ecology,Evolution and Systematics, 17 (3):243-254.
[25]	Pu Xiao-hong, Yue Xiu-le, An Li-zhe. 2017. The response of plant to UV-B radiation. Scientia Sinica Vitae, 47 (8):818-828. (in Chinese)
	蒲晓宏, 岳修乐, 安黎哲. 2017. 植物对UV-B辐射的响应与调控机制. 中国科学:生命科学, 47 (8):818-828.
[26]	Schettini E, de Salvador F R, Scarascia-Mugnozza G, Vox G. 2011. Radiometric properties of photoselective and photoluminescent greenhouse plastic films and their effects on peach and cherry tree growth. The Journal of Horticultural Science and Biotechnology, 86 (1):79-83.
[27]	Shao Mao-ni. 2017. Study on nano light-transfer membrane for facility nectarine[M. D. Dissertation]. Nanjing: Nanjing Normal University. (in Chinese)
	邵毛妮. 2017. 设施油桃专用纳米转光膜的研究[硕士论文]. 南京: 南京师范大学.
[28]	Wang Feng, Yan Jia-rong, Chen Xue-yu, Jiang Cheng-hao, Meng Si-da, Liu Yu-feng, Xu Tao, Qi Ming-fang, Li Tian-lai. 2019. Light regulation of chlorophyll biosynthesis in plants. Acta Horticulturae Sinica, 46 (5):975-994. (in Chinese) doi: 10.16420/j.issn.0513-353x.2018-0799
	王峰, 闫家榕, 陈雪玉, 姜程浩, 孟思达, 刘玉凤, 许涛, 齐明芳, 李天来. 2019. 光调控植物叶绿素生物合成的研究进展. 园艺学报, 46 (5):975-994. doi: 10.16420/j.issn.0513-353x.2018-0799
[29]	Wang Hong, Yue Kun, Yang Chengkun, Guo Yujian, Wang Peixuan, Zhou Kaibing. 2020. Physiological reasons for inhibiting photosynthesis of mango leaves by enhanced UV-B radiation. Acta Horticulturae Sinica, 47 (2):242-252. (in Chinese) doi: 10.16420/j.issn.0513-353x.2019-0381
	王红, 岳堃, 杨成坤, 郭钰柬, 王珮璇, 周开兵. 2020. UV-B辐射增强处理抑制杧果叶片光合作用的生理原因分析. 园艺学报, 47 (2):242-252. doi: 10.16420/j.issn.0513-353x.2019-0381
[30]	Wang Jun. 2004. Study on application of UV-B supplement combined with CO₂ enrichment in winter plastic greenhouse tomato planting[M. D. Dissertation]. Yangling: Northwest A & F University. (in Chinese)
	王军. 2004. 补充紫外UV-B辐射和CO₂施肥在冬季大棚番茄生产中的应用研究[硕士论文]. 杨凌: 西北农林科技大学.
[31]	Wang Ling, Deng Lin-ying, Li Xi. 2014. Effects of SO₂ stress on chlorophyll synthesis and metabolism in golden leaf plants. Chinese Journal of Ecology, 33 (9):2382-2387. (in Chinese)
	王玲, 邓林颖, 李西. 2014. SO₂胁迫对金色叶植物叶片叶绿素合成代谢的影响. 生态学杂志, 33 (9):2382-2387.
[32]	Wang P, Grimm B. 2021. Connecting chlorophyll metabolism with accumulation of the photosynthetic apparatus. Trends in Plant Science, 26 (5):484-495. doi: 10.1016/j.tplants.2020.12.005 pmid: 33422426
[33]	Wang Xiu-pin, Li Xiang, Li Zu-ran, Ling Cheng-ting, He Yong-mei, Li Yuan. 2022. Research progress on the characteristics and mechanisms of transgenerational plasticity of plants in response to enhanced UV-B radiation. Plant Physiology Journal, 58 (5):797-805. (in Chinese)
	王修蘋, 李想, 李祖然, 凌成婷, 何永美, 李元. 2022. 植物对UV-B辐射增强响应的跨代可塑性特征及机制研究进展. 植物生理学报, 58 (5):797-805.
[34]	Xiao Wei,Wu Hong-yu,Li Ling,Chen Xiu-de,Li Dong-mei,Liu Li-chang,Gao Dong-sheng. 2016. Facilities cultivation of peach,apricot,plum special tree thin column(a stick),plastic technology. China Fruits,(2):78-80,103. (in Chinese)
	肖伟, 武红玉, 李玲, 陈修德, 李冬梅, 刘立常, 高东升. 2016. 设施栽培桃、杏、李专用树形细柱形(一根棍)整形技术. 中国果树,(2):78-80,103.
[35]	Xie L, Solhaug K A, Song Y, Johnsen B, Tollefsen K E. 2020. Effects of artificial ultraviolet B radiation on the macrophyte Lemna minor:a conceptual study for toxicity pathway characterization. Planta, 252 (5):86. doi: 10.1007/s00425-020-03482-3 pmid: 33057834
[36]	Yang Jing, Zhang Guang-qun, Li Ming-rui, Zhan Fang-dong, Li Yuan, Zu Yan-qun, He Yong-mei. 2022. Effects of UV-B radiation and nitrogen application on the growth,mineral nutrition,and antioxidant physiology of Azolla. Journal of Agricultural Resources and Environment, 39 (3):567-574. (in Chinese)
	杨竞, 张光群, 李明锐, 湛方栋, 李元, 祖艳群, 何永美. 2022. UV-B辐射和施氮对满江红生长、矿质营养和抗氧化生理的影响. 农业资源与环境学报, 39 (3):567-574.
[37]	Yang Qing, Aishajiang Mai-mai-ti, Wang Zhi-xia, Liu Guo-jie. 2012. Effects of DA-6 on chlorophyll biosythesis pathway in peach leaves. Acta Horticulturae Sinica, 39 (4):621-628. (in Chinese)
	杨清,艾沙江 · 买买提,王志霞,刘国杰. 2012. DA-6对桃树叶片叶绿素合成途径的调控研究. 园艺学报, 39 (4):621-628.
[38]	Yu Min,Hu Cheng-xiao,Wang Yun-hua. 2006. Effects of molybdenum on the precursors of chlorophyll biosynthesis in winter wheat cultivars under low temperature. Scientia Agricultura Sinica,(4):702-708. (in Chinese)
	喻敏, 胡承孝, 王运华. 2006. 低温条件下钼对冬小麦叶绿素合成前体的影响. 中国农业科学,(4):702-708.
[39]	Yu Ni-na, Li Dong-mei, Tan Qiu-ping, Zhang Hai-sen, Gao Dong-sheng. 2013. Effect of UV-B radiation on assimilate translocation and distribution in fruiting shoot of protected peach. Chinese Journal of Applied & Environmental Biology, 19 (1):157-163. (in Chinese)
	于妮娜, 李冬梅, 谭秋平, 张海森, 高东升. 2013. UV-B辐射对设施桃结果枝同化物转运和分配的影响. 应用与环境生物学报, 19 (1):157-163.
[40]	Zhang Ke-kun, Liu Feng-zhi, Wang Xiao-di, Shi Xiang-bin, Wang Bao-liang, Zheng Xiao-cui, Ji Xiao-hao, Wang Hai-bo. 2017. Effects of supplementary light with different wavelengths on fruit quality of‘Ruidu Xiangyu’grape under promoted cultivation. Chinese Journal of Applied Ecology, 28 (1):115-126. (in Chinese)
	张克坤, 刘凤之, 王孝娣, 史祥宾, 王宝亮, 郑晓翠, 冀晓昊, 王海波. 2017. 不同光质补光对促早栽培‘瑞都香玉’葡萄果实品质的影响. 应用生态学报, 28 (1):115-126. doi: 10.13287/j.1001-9332.201701.003
[41]	Zhang Xiu-de. 2016. Gene cloning and functional identification of MdHEMA1encoding the key enzyme for chlorophyll biosynthesis in apple[M. D. Dissertation]. Beijing: Chinese Academy of Agricultural Sciences. (in Chinese)
	张修德. 2016. 苹果叶绿素合成关键酶基因MdHEMA1克隆和功能验证[硕士论文]. 北京: 中国农业科学院.
[42]	Zhao Shi-jie, Cang Jing. 2015. Experimental guidelines in plant physiology. Beijing: China Agriculture Press. (in Chinese)
	赵世杰, 苍晶. 2015. 植物生理学实验指导. 北京: 中国农业出版社.
[43]	Zheng Fu-xing, Yan An, Gao Xue, Yan Yong-liang, Wang Rui, Geng Hong-wei. 2021. Genome-wide association scanning of chlorophyll SPAD in wheat under water and drought treatments. Journal of Plant Genetic Resources, 22 (5):1334-1347. (in Chinese)
	郑福兴, 颜安, 高雪, 严勇亮, 王睿, 耿洪伟. 2021. 水旱处理下小麦叶绿素相对含量全基因组关联分析. 植物遗传资源学报, 22 (5):1334-1347. doi: 10.13430/j.cnki.jpgr. 20210309003
[44]	Zhou Xin-ming. 2007. The effects of enhanced Ultraviolet-B radiation on photosynthetic physiology and total hydroxybenzene in the leaves of grapevine[M. D. Dissertation]. Yangling: Northwest A & F University. (in Chinese)
	周新明. 2007. UV-B辐射增强对葡萄叶片光合生理及总酚含量的影响[硕士论文]. 杨凌: 西北农林科技大学.
[45]	Zhu Xue-yun, Li Jian-mou, Wang Hai-yan, Kong Fan-qin. 2017. Analysis of chlorophyll biosynthesis and chloroplast microstructure in Euonymus japonicus. var. aureamarginatus and E. japonicus var. aureovariegatus. Southwest China Journal of Agricultural Sciences, 30 (8):1767-1771. (in Chinese)
	朱雪云, 李建谋, 王海燕, 孔凡芹. 2017. 金边黄杨和金心黄杨叶绿素合成与叶绿体结构分析. 西南农业学报, 30 (8):1767-1771.