Advances of Light-harvesting Chlorophyll a/b Binding Protein (CAB) in Higher Plants
ZHONG Si-Tong1,2, ZHANG Ya-Zhen2, KONG Xiang-Rui2, JIN Shan1, CHEN Zhi-Hui2, CHEN Chang-Song1,2,3,*
1 College of Horticulture, Fujian Agriculture and Forestry University, Fuzhou 350002, China; 2 Tea Research Institute, Fujian Academy of Agricultural Sciences/Fujian Branch, National Center for Tea Improvement, Fuzhou 350012, China; 3 National Agricultural Experimental Station for Soil Quality, Fu'an 355015, China
Abstract:Light-harvesting chlorophyll a/b binding protein (CAB) is a kind of membrane protein encoded by the CAB gene family. It plays an important role in capturing and transmitting light energy, photoprotection and excess energy dissipation, regulating the distribution of light energy between the two photosystems, and maintaining the structure of the thylakoid membrane. CAB genes widely participate in the whole biological processes in higher plants, including the growth and development of leaf, flower and fruit, and in response to various stresses. In this review, the research progresses of CAB family in their classification, protein structure, physiological functions, and stress expression characteristics were summarized. The future research directions of CAB were also prospected. This paper provides references for further studies on the CAB gene family.
[1] 陈勇智, 苏良辰, 李玲. 2018. TSA对不同抗旱性花生品种响应干旱过程中光合特性及相关基因表达的影响[J]. 华南师范大学学报:自然科学版, 50(3): 72-77. (Chen Z Y, Su L C, Li L, et al.2018. Effect of TSA on photosynthesis-related gene expression in different drought-resistant peanut cultivars subjected to drought stress[J]. Journal of South China Normal University (Natural Science Edition), 50(3): 72-77.) [2] 丁凯旋, 郑婉茹, 李琳琳, 等. 2021. 木薯MeLHCB4基因的克隆及表达分析[J]. 热带作物学报, 42(10): 2813-2818. (Ding K X, Zheng W R, Li L L, et al.2021. Cloning and expression analysis of MeLHCB4 from Cassava[J]. Chinese Journal of Tropical Crops, 42(10): 2813-2818.) [3] 高志民, 刘颖丽, 彭镇华. 2012. 毛竹PSII大量捕光天线蛋白基因克隆及其表达分析[J]. 植物科学学报, 30(1): 64-71. (Gao Z M, Liu Y L, Peng Z H.2012. Cloning and molecular characterization of the major light harvesting antenna protein genes of PSⅡ in Phyllostachys edulis[J]. Plant Science Journal, 30(1): 64-71.) [4] 葛宇, 董相书, 吴斌, 等. 2018. 油梨果肉捕光叶绿素a/b结合蛋白基因PaCAB1的克隆、序列分析及表达[J]. 中国南方果树, 47(05): 18-22. (Ge Y, Dong X S, Wu B, et al.2018. Cloning, sequence analysis and expression of chlorophyll a/b binding protein PaCAB1 from Persea americana[J]. South China Fruits, 47(05): 18-22.) [5] 韩霜, 刘瑞霞, 张兆和, 等. 2013. 菊花叶绿素a/b结合蛋白基因CmLhcb1及其启动子的克隆和表达分析[J]. 园艺学报, 40(06): 1119-1128. (Han S, Liu R X, Zhang Z H, et al.2013. Cloning of chlorophyll a/b binding protein CmLhcb1 and promoter from Chrysanthemum morifolium and expression analysis[J]. Acta Horticulturae Sinica, 40(06): 1119-1128.) [6] 郝利民, 王洪亮, 梁厚果. 1999. 复水对玉米光系统II捕光叶绿素a/b蛋白复合体的影响[J]. 植物学报, 41(06): 613-616. (Hao L M, Wang H L, Liang H G, et al.1999. Effects of rewatering on light-harvesting chlorophyll a/b protein complex of photosystem II in Zea mays[J]. Acta Botanica Sinica, 41(06): 613-616.) [7] 贺军虎, 李唯正, 陈华蕊, 等. 2017. '金煌'杧果MiCAB2的克隆及表达与花期调控关系分析[J]. 园艺学报, 44(7): 1275-1286. (He J H, Li W Z, Chen H R, et al.2017. Cloning of the light harvesting chlorophyll a/b binding protein gene (MiCAB2) from 'Jinhuang' Mango, and correlation analysis between its expression and flowering regulation[J]. Acta Horticulturae Sinica, 44(7): 1275-1286.) [8] 胡志航, 金伟刚, 宋丽双, 等. 2020. 水稻编码光系统Ⅱ相关捕光色素蛋白Lhcb2基因的克隆与表达分析[J]. 扬州大学学报:农业与生命科学版, 41(4): 1-8. (Hu Z H, Jin W G, Song L S, et al.2020. Cloning and expression analysis of the Lhcb2 gene encoding light-harvesting complex protein in rice[J]. Journal of Yangzhou University: Agricultural and Life Science Edition, 41(4):1-8.) [9] 黄春雪. 2022. 甜菜叶绿素a-b结合蛋白基因BvLhcb-2的克隆与表达特性初步分析[D]. 硕士学位论文, 黑龙江大学, 导师: 王希, pp. 28-41. (Huang C X.2022. Cloning and expression analysis of chlorophyll a/b binding protein gene BvLhcb-2 in Beta vulgaris[D]. Thesis for M.S., Heilongjiang University, Supervisor: Wang X, pp. 28-41.) [10] 黄巍, 李静, 庄艺, 等. 2022. 植物生物钟抵御生物胁迫研究进展[J]. 华南农业大学学报, 43(06): 121-129. (Huang W, Li J, Zhuang Y, et al.2022. Timing of plant immune responses to biotic stress[J]. Journal of South China Agricultural University, 43(06): 121-129) [11] 蒋向辉, 苑静. 2016. 金银花叶绿素a/b结合蛋白基因LjCab克隆与表达特性分析[J]. 华中师范大学学报(自然科学版), 50(03): 409-414. (Jiang X H, Yuan J.2016. Cloning and expression characterization of LjCab gene of the light harvesting chlorophyll a/b binding protein from Lonicera japonica Thunb[J].Journal of Central China Normal University, 50(03): 409-414.) [12] 李利超, 孙化雨, 娄永峰, 等. 2015. 毛竹捕光色素结合蛋白基因结构及表达模式分析[J]. 世界竹藤通讯, 13(06): 1-8. (Li L C, Sun H Y, Lou Y F, et al.2015. Structure and expression analysis of light-harvesting chlorophyll a/b binding protein genes in moso bamboo (Phyllostachys edulis)[J]. World Bamboo And Rattan, 13(06): 1-8.) [13] 李玲, 曲嘉琪, 时丕秀, 等. 2021. 基于转录组测序的薰衣草响应温度变化的光合作用分子机制[J/OL].分子植物育种:1-14. (Li L, Qu J Q, Shi P X, et al.2021. Molecular mechanism of photosynthesis in response to temperature change based on transcriptome sequencing in Lavandula angustifolia Mill[J/OL]. Molecular Plant Breeding: 1-14) [14] 李韵佳. 2021. 低温胁迫下三倍体枇杷微量捕光天线蛋白基因EjLhcb4.1/5/6的功能分析[D]. 硕士学位论文, 西南大学, 导师: 郭启高, pp. 61-68. (Li Y J.2021. Functional characterization of light harvesting chlorophyll a/b binding antenna proteins EjLhcb4.1/5/6 genes in triploid loquat (Eriobotrya japonica) under cold stress[D]. Thesis for M.S., Southwest University, Supervisor: Guo Q G, pp. 61-68.) [15] 李真, 刘明英, 韩小娇, 等. 2014. 东南景天捕光叶绿素a/b结合蛋白基因SaLhcb2的分离及功能[J]. 浙江农林大学学报, 31(6): 838-846. (Li Z, Liu M Y, Han X Y, et al.2014. Characterization of a light-harvesting chlorophyll a/b binding protein (LHCB) gene, SaLhcb2, in Sedum alfredii[J]. Journal of Zhejiang A&F University, 31(6): 838-846.) [16] 李祖任, 罗丁峰, 柏浩东,等. 2021. 小飞蓬捕光叶绿素结合蛋白基因CcLhca-J9克隆及表达分析[J]. 中国农业科学, 54(1): 86-94. (Li Z R, Luo D F, Bai H D, et al.2021. Cloning and expression analysis of light harvesting chlorophyll a/b protein gene CcLhca-J9 in Conyza canadensis[J]. Scientia Agricultura Sinica, 54(1): 86-94.) [17] 林江波, 王伟英, 邹晖,等. 2013. 中国水仙叶绿素a/b结合蛋白基因Ntcab1的克隆与序列分析[J]. 福建农业学报, 28(5): 463-467. (Lin J B, Wang W Y, Zou H, et al.2013. Cloning and sequence analysis and gene of chlorophyll a/b binding protein from Narcissus tazettz var. chinensis[J]. Fujian Journal of Agricultural Sciences, 28(5): 463-467.) [18] 刘建汀, 谢贵水, 莫业勇, 等. 2014. 橡胶树捕光叶绿素结合蛋白基因Lhcb2.2的克隆与分析[J]. 西南农业学报, 27(06): 2267-2271. (Liu J T, Xie G S, Mo Y Y, et al.2014. Molecular cloning and expression analysis of HbLhcb2.2 gene from rubber tree (Hevea brasiliensis Muell. Arg.)[J]. Southwest China Journal of Agricultural Sciences, 27(06): 2267-2271.) [19] 刘亚丽, 张凯敏, 高彩球. 2014. 刚毛柽柳光捕获叶绿素a/b结合蛋白基因Thcab1的克隆与分析[J]. 东北林业大学学报, 42(02): 52-56. (Liu Y L, Zhang K M, Gao C Q, et al.2014. Chlorophyll a/b binding proteins Thcab1 gene clone and expression analysis of Tamarix hispida[J]. Journal of Northeast Forestry University, 42(02): 52-56.) [20] 罗玲. 2008. 捕光色素蛋白复合物的研究进展[J]. 现代农业科技, (22): 270-273. (Luo L. 2008. Research progress of light-harvesting pigment protein complexes[J]. Modern Agricultural Science and Technology, (22): 270-273.) [21] 梅菊芬, 徐德良, 汤茶琴, 等. 2018. 茶树CsLhcb4基因的电子克隆与生物信息学分析[J]. 江苏农业科学, 46(09): 51-55. (Mei J F, Xu D L, Tang C Q, et al.2018. Electronic cloning and bioinformatics analysis of CsLhcb4 gene in Camellia sinensis[J]. Jiangsu Agricultural Sciences , 46(09): 51-55.) [22] 石兰馨, 张晓平, 梁厚果. 1995. 捕光叶绿素a/b结合蛋白和cab基因[J]. 植物生理学通讯, 31(06): 470-476. (Shi L X, Zhang X P, Liang H G.1995. Light harvesting chlorophyll a/b binding protein and cab gene[J]. Plant Physiology Communications, 31(06): 470-476.) [23] 宋丽莉, 吴琼, 吉喜燕, 等. 2022. 高光胁迫下香果树叶片生理生化指标变化及相关基因差异表达和功能分析[J]. 植物资源与环境学报, 31(06): 52-62;83.(Song L L, Wu Q, Ji X Y, et al. 2022. Variations of physiological and biochemical indexes and differential expression and functional analyses on related genes in leaves of Enmenopferys henryi under high light stress[J]. Journal of Plant Resources and Environment, 31(06): 52-62;83.) [24] 孙钦秒, 冷静, 李良璧, 等. 2000a. 高等植物光系统II捕光色素蛋白复合体结构与功能研究的新进展[J] .植物学通报, 17(04): 289-301. (Sun Q M, Leng J, Li L B, et al.2000a. Recent advances of studies on the structure and function of the light-harvesting chlorophyll a/b protein complex[J]. Chinese Bulletin of Botany, 17(04): 289-301.) [25] 孙钦秒, 李良璧, 阎久胜, 等. 2000b. 光照和温度对豌豆Lhcb2基因表达的影响(英文)[J]. 植物学报, 42(03): 258-262. (Sun Q M, Li L B, Yan J S, et al.2000b. Effects of light and temperature on the expression of the Lhcb2 gene in pea[J]. Acta Botanica Sinica, 42(03): 258-262.) [26] 孙世磊. 2022. 玉米光合相关转录因子的鉴定、突变体创制及功能分析[D].博士学位论文, 山东农业大学,导师: 李平华, pp. 73-77. (Sun S L.2022. Identification, mutant creation, and functional analysis of photosynthesis-related transcription factors in maize[D]. Thesis for Ph. D., Shandong Agricultural University, Supervisor: Li P H, pp. 73-77.) [27] 唐文莉. 2008. 毛竹Lhca基因的克隆和光照在转录水平对其表达的调控[D]. 博士学位论文, 中国林业科学研究院, 导师: 彭镇华, pp. 82-85. (Tang W L.2008. Cloning Lhca genes and at the transcriptional level light intensity regulating their expression of Phyllostachys edulis[D]. Thesis for Ph. D., Chinese Academy of Forestry, Supervisor: Peng Z H, pp. 82-85.) [28] 田月月. 2020. 黄金芽茶树叶色响应光质的生理特性及机制研究[D]. 博士学位论文, 山东农业大学, 导师: 张丽霞, pp. 58-69. (Tian Y Y.2020. Mechanism of physiological characteristics of leaf color in Camellia sinensis cv. Huangjinya response to light quality[D]. Thesis for Ph. D., Shandong Agricultural University, Supervisor: Zhang L X, pp. 58-69.) [29] 王小佳. 2020. 雷波脐橙及其早熟单株果实脱绿期的叶绿素降解代谢关键基因表达分析[D]. 硕士学位论文, 四川农业大学, 导师: 汪志辉, pp.36-40. (Wang X J.2020. Differential expression analysis of key genes for chlorophyll degradation and metabolism of Leibo navel orange and its early-ripening fruit mutation[D]. Thesis for M.S., Sichuan Agricultural University, Supervisor: Wang Z H, pp. 36-40.) [30] 王云鹤. 2020. 苎麻Lhc基因的鉴定和表达模式研究[D]. 硕士学位论文, 华中农业大学, 导师: 彭定祥, pp. 55-61. (Wang Y H.2020. Identification and expression profiling of Lhc genes in ramie[D]. Thesis for M.S., Huazhong Agricultural University, Supervisor: Peng D X, pp. 55-61.) [31] 未丽, 徐秉良, 雷江丽, 等. 2008. 胡杨叶绿素a/b结合蛋白基因的克隆及序列特性分析[J].中国农业科技导报, (04): 63-69. (Wei L, Xu B L, Lei J L, et al.2008. Cloning and characterization of a full-length cab gene encoding the light harvesting chlorophyll a/b binding protein from Populus euphratica[J]. Journal of Agricultural Science and Technology, 2008(04): 63-69.) [32] 吴凤华. 2022. 过表达拟南芥Lhca5对其PSI-LHCI结构及光谱学特征的影响[D]. 硕士学位论文, 济南大学, 导师: 秦晓春, pp. 61-64. (Wu F H.2022. Effect of overexpression of Arabidopsis thaliana Lhca5 on its PSI-LHCI structural and spectroscopic features[D]. Thesis for M.S., University of Jinan, Supervisor: Qin X C, pp. 61-64.) [33] 阳江华, 张希财, 邹智. 2019. 橡胶树捕光叶绿素a/b结合蛋白基因CAB2的克隆与分析[J].西南林业大学学报(自然科学), 39(01): 88-94. (Yang J H, Zhang X C, Zou Z.2019. Molecular cloning and analysis of HbCAB2, A chlorophylla/b-binding protein-encoding gene from Hevea brasiliensis[J]. Journal of Southwest Forestry University, 39(01): 88-94.) [34] 翟玉山, 邓宇晴, 董萌, 等. 2016. 甘蔗捕光叶绿素a/b结合蛋白基因ScLhca3的克隆及表达[J]. 作物学报, 42(09): 1332-1341. (Zhai Y S, Deng Y Q, Dong M, et al.2016. Cloning and characterization of light harvesting chlorophyll a/b binding protein coding gene (ScLhca3) in sugarcane[J]. Acta Agronomica Sinica, 42(09): 1332-1341.) [35] 曾水玉. 2012. 万寿菊试管开花及其分子机制的研究[D]. 硕士学位论文, 福建农林大学, 导师: 赖钟雄, pp. 72-77. (Zeng S Y.2012. Studies on in vitro flowering and related molecular mechanism in Tagetes erecta L.[D]. Thesis for M.S., Fujian Agricultural University, Supervisor: Lai Z X, pp. 72-77.) [36] 张计育, 潘德林, 贾展慧, 等. 2018. 中华猕猴桃品种'Hort16A'果肉颜色形成的分子机制[J]. 植物资源与环境学报, 27(03): 1-10. (Zhang J Y, Pan D L, Jia Z H, et al.2018. Molecular mechanism of flesh color formation of Actinidia chinensis 'Hort16A'[J]. Journal of Plant Resources and Environment, 27(03): 1-10.) [37] 张蕾, 于永昂. 2020. 小麦TaLhca基因的克隆及表达分析[J]. 西北植物学报, 40(03): 394-402. (Zhang L, Yu Y A.2020. Cloning and expression analysis of TaLhca gene in wheat[J]. Acta Botanica Boreali-Occidentalia Sinica, 40(03): 394-402.) [38] 张帅, 崔金杰, 吕丽敏, 等. 2012. 棉花3种抗逆相关基因对苗蚜为害的应答反应[J].棉花学报, 24(03): 215-221. (Zhang S, Cui J J, Lu L L, et al.2012. Response of three resistance genes to damage caused by Aphis gossypii (Hemiptera: Aphididae) in cotton[J].Cotton Science, 24(03): 215-221.) [39] 张亚杰. 2006. 高等植物光系统II大量捕光色素蛋白复合体的稳定性研究[D]. 博士学位论文,中国科学院研究生院(植物研究所), 导师: 杨春虹, pp. 94-98. (Zhang Y J.2006. Stability of the major light-harvesting complex of photosystem II of higher plant[D]. Thesis for Ph. D., Chinese Academy of Sciences, Supervisor: Yang C H, pp. 94-98.) [40] 赵鹏飞. 2019. 低温胁迫下菊花叶片光系统特性等生理指标分析及相关基因表达[D]. 硕士学位论文, 河南农业大学, 导师: 李永华, pp. 36-39. (Zhao P F.2019. Analysis of photosystem characteristics, physiological indicators and expression of related genes under low temperature stress in Chrysanthemum leaves[D]. Thesis for M.S., Henan Agricultural University, Supervisor: Li Y H, pp. 36-39.) [41] 赵奇, 茹京娜, 李宜统, 等. 2022. 小麦Lhc基因家族鉴定与表达模式分析[J]. 植物遗传资源学报, 23(06): 1766-1781. (Zhao Q, Ru J N, Li Y T, et al.2022. Identification and expression pattern analysis of Lhc gene family members in wheat[J]. Journal of Plant Genetic Resources, 23(06): 1766-1781.) [42] 赵彦, 高鑫, 王丹, 等. 2017. 蒙古冰草Lhcb1基因克隆及干旱胁迫下的表达分析[J]. 西北植物学报, 37(02): 211-216. (Zhao Y, Gao X, Wang D, et al.2017. Cloning and expression analysis of Lhcb1 from Agropyron mongolicum under drought stress[J]. Acta Botanica Boreali-Occidentalia Sinica, 37(02): 211-216.) [43] 朱彬彬, 崔百明, 向本春. 2017. 番茄叶绿素a/b结合蛋白基因cab-1a的克隆与定位[J]. 江苏农业科学, 45(14): 20-23. (Zhu B B, Cui B M, Xiang B C.2017. Cloning and mapping of chlorophyll a/b binding protein gene cab-1a from tomato[J]. Jiangsu Agricultural Sciences, 45(14): 20-23.) [44] 邹寒艳. 2015. 水稻OsLIR1维持叶绿体功能的机制及OsEXPB2参与水稻根系发育的功能研究[D]. 硕士学位论文, 重庆大学, 导师: 王贵学, pp. 81-83. (Zou H Y.2015. Study on mechanisms of OsLIR1 maintaining chloroplast function and roles of OsEXPB2 in root development in rice (Oryza sativa)[D]. Thesis for M.S., Chongqing University, Supervisor: Wang G X, pp. 81-83.) [45] 邹智, 杨礼富, 安峰, 等. 2013. 橡胶树AtCAB1同源基因的克隆及其在稳定与衰老期叶片中的差异分析[J]. 热带农业科学, 33(4): 30-35. (Zou Z, Yang L F, An F, et al.2013. Molecular cloning of a cDNA homologue to Arabidopsis CAB1 from Hevea brasiliensis and its different expression in mature and senescent rubber tree leaves[J]. Chinese Journal of Tropical Agriculture, 33(4): 30-35.) [46] Andersson J, Wentworth M, Walters R G, et al.2003. Absence of the lhcb1 and lhcb2 proteins of the light-harvesting complex of photosystem II - effects on photosynthesis, grana stacking and fitness[J]. The Plant Journal, 35(3): 350-361. [47] Ballottari M, Dall'Osto L, Morosinotto T, et al.2007. Contrasting behavior of higher plant photosystem I and II antenna systems during acclimation[J]. Journal of Biological Chemistry, 282(12): 8947-8958. [48] Bartholomew D M, Bartley G E, Scolnik P A.1991. Abscisic acid control of rbcS and cab transcription in tomato leaves[J]. Plant Physiology, 96(1): 291-296. [49] Bassett C L, Callahan A M, Artlip T S, et al.2007. A minimal peach type II chlorophyll a/b binding protein promoter retains tissue-specificity and light regulation in tomato[J]. BMC Biotechnology, 7(1): 47. [50] Bian Z, Yang Q, Li T, et al.2018. Study of the beneficial effects of green light on lettuce grown under short‐term continuous red and blue light‐emitting diodes[J]. Physiologia Plantarum, 164(2): 226-240. [51] Büchel C.2015. Evolution and function of light harvesting proteins[J]. Journal of Plant Physiology, 172: 62-75. [52] Cannon S B, Mitra A, Baumgarten A, et al.2004. The roles of segmental and tandem gene duplication in the evolution of large gene families in Arabidopsis thaliana[J]. BMC Plant Biology, 4(10): 1-21. [53] Chang Y C, Walling L L.1991. Abscisic acid negatively regulates expression of chlorophyll a/b binding protein genes during soybean embryogeny[J]. Plant Physiology, 97(3): 1260-1264. [54] Chen X F, Li J J, Yu Y, et al.2022. STAY-GREEN and light-harvesting complex II chlorophyll a/b binding protein are involved in albinism of a novel albino tea germplasm 'Huabai 1'[J]. Scientia Horticulturae, 293: 110653. [55] Christensen S, Silverthorne J.2001. Origins of phytochrome-modulated Lhcb mRNA expression in seed plants[J]. Plant physiology, 126(4): 1609-1618. [56] Churin Y, Adam E, Kozma-Bognar L, et al.2003. Characterization of two Myb-like transcription factors binding to CAB promoters in wheat and barley[J]. Plant Molecular Biology, 52(2): 447-462. [57] Damkjaer J T, Kereiche S, Johnson M P, et al.2009. The photosystem II light-harvesting protein Lhcb3 affects the macrostructure of photosystem II and the rate of state transitions in Arabidopsis[J]. The Plant Cell, 21(10): 3245-3256. [58] De Bianchi S, Betterle N, Kouril R, et al.2011. Arabidopsis mutants deleted in the light-harvesting protein lhcb4 have a disrupted photosystem II macrostructure and are defective in photoprotection[J]. The Plant Cell, 23(7): 2659-2679. [59] De Bianchi S, Dall'Osto L, Tognon G, et al.2008. Minor antenna proteins CP24 and CP26 affect the interactions between photosystem II subunits and the electron transport rate in grana membranes of Arabidopsis[J]. Plant Cell, 20(4): 1012-1028. [60] Deng Y S, Kong F Y, Zhou B, et al.2014. Heterology expression of the tomato LeLhcb2 gene confers elevated tolerance to chilling stress in transgenic tobacco[J]. Plant Physiology and Biochemistry, 80: 318-327. [61] Dunsmuir P.1985. The petunia chlorophyll a/b binding protein genes: A comparison of Cab genes from different gene families[J]. Nucleic Acids Research, 13(7): 2503-2518. [62] Huang J, Zhao X, Chory J.2019. The Arabidopsis transcriptome responds specifically and dynamically to high light stress[J]. Cell Reports, 29(12): 4186-4199. [63] Horton P, Johnson M P, Perez-Bueno M L, et al.2008. Photosynthetic acclimation: Does the dynamic structure and macro-organization of photosystem II in higher plant grana membranes regulate light harvesting states?[J]. The FEBS Journal, 275(6): 1069-1079. [64] Jansson S, Pichersky E, Bassi R, et al.1992. A nomenclature for genes encoding the chlorophyll a/b binding proteins of higher plants[J]. Plant Molecular Biology Reporter, 10(3): 242-253. [65] Jensen P E, Bassi R, Boekema E J, et al.2007. Structure, function and regulation of plant photosystem I[J]. Biochimica et Biophysica Acta, 1767(5): 335-352. [66] Jiang Q, Xu Z S, Wang F, et al.2014. Effects of abiotic stresses on the expression of Lhcb1 gene and photosynthesis of Oenanthe javanica and Apium graveolens[J]. Biologia Plantarum, 58(2): 256-264. [67] Klimmek F, Sjödin A, Noutsos C, et al.2006. Abundantly and rarely expressed Lhc protein genes exhibit distinct regulation patterns in plants[J]. Plant Physiology, 140(3): 793-804. [68] Labate M T, Ko K, Ko Z W, et al.2004. Constitutive expression of pea Lhcb1-2 in tobacco affects plant development, morphology and photosynthetic capacity[J]. Plant Molecular Biology, 55(5): 701-714. [69] Li X W, Zhu Y L, Chen C Y, et al.2020. Cloning and characterization of two chlorophyll a/b binding protein genes and analysis of their gene family in Camellia sinensis[J]. Scientific Reports, 10(1): 4602-4633. [70] Lin W, Guo X, Pan X, et al.2018. Chlorophyll composition, chlorophyll fluorescence, and grain yield change in esl mutant rice[J]. International Journal of Molecular Sciences, 19(10): 2945. [71] Liu M, Zhang S, Hu J, et al.2019. Phosphorylation-guarded light-harvesting complex II contributes to broad-spectrum blast resistance in rice[J]. Proceedings of the National Academy of Sciences of the USA, 116(35): 17572-17577. [72] Liu R, Xu Y H, Jiang S C, et al.2013. Light-harvesting chlorophyll a/b binding proteins, positively involved in abscisic acid signaling, require a transcription repressor, WRKY40, to balance their function[J]. Journal of Experimental Botany, 64(18): 5443-5456. [73] Luo J, Abid M, Tu J, et al.2022. Genome-wide identification of the LHC gene family in kiwifruit and regulatory role of AcLhcb3.1/3.2 for chlorophyll a content[J]. International Journal of Molecular Sciences, 23(12): 6528. [74] Ma Q P, Li H, Zou Z W, et al.2018. Transcriptomic analyses identify albino-associated genes of a novel albino tea germplasm 'Huabai 1'[J]. Horticulture Research, 5(1): 123-134. [75] Maïna F, Bassi R, Robaglia C, et al.2013. Post-transcriptional control of light-harvesting genes expression under light stress[J]. Plant Molecular Biology, 82(1-2): 147-154. [76] Neilson J A D, Durnford D G.2010. Structural and functional diversification of the light-harvesting complexes in photosynthetic eukaryotes[J]. Photosynthesis Research, 106(1): 57-71. [77] Neoh B K, Wong Y C, Teh H F, et al.2019. Diurnal biomarkers reveal key photosynthetic genes associated with increased oil palm yield[J]. PLOS ONE, 14(3): e0213591. [78] Oberschmidt O, Hücking C, Piechulla B.1995. Diurnal Lhc gene expression is present in many but not all species of the plant kingdom[J]. Plant Molecular Biology, 27: 147-153. [79] Pan X, Liu Z, Li M, et al.2013. Architecture and function of plant light-harvesting complexes II[J]. Current Opinion in Structural Biology, 23(4): 515-525. [80] Perveen S, Qu M, Chen F, et al.2020. Overexpression of maize transcription factor mEmBP-1 increases photosynthesis, biomass, and yield in rice[J]. Journal of Experimental Botany, 71(16): 4944-4957. [81] Pietrzykowska M, Suorsa M, Semchonok D A, et al.2014. The light-harvesting chlorophyll a/b binding proteins Lhcb1 and Lhcb2 play complementary roles during state transitions in Arabidopsis[J]. The Plant Cell, 26(9): 3646-3660. [82] Qiao G, Wen X P, Zhang T.2015. Molecular cloning and characterization of the light-harvesting chlorophyll a/b gene from the pigeon pea (Cajanus cajan)[J]. Applied Biochemistry and Biotechnology, 177(7): 1447-1455. [83] Ruban A V, Wentworth M, Yakushevska A E, et al.2003. Plants lacking the main light-harvesting complex retain photosystem II macro-organization[J]. Nature, 421(6923): 648-652. [84] Sakuraba Y, Schelbert S, Park S Y, et al.2012. STAY-GREEN and chlorophyll catabolic enzymes interact at light-harvesting complex II for chlorophyll detoxification during leaf senescence in Arabidopsis[J]. Plant Cell, 24(2): 507-518. [85] Sato R, Ito H, Tanaka A.2015. Chlorophyll b degradation by chlorophyll b reductase under high-light conditions[J]. Photosynthesis Research, 126(2-3): 249-259. [86] Sheen J Y, Bogorad L.1986. Differential expression of six light-harvesting chlorophyll a/b binding protein genes in maize leaf cell types[J]. Proceedings of the National Academy of Sciences of the USA, 83(20): 7811-7815. [87] Staneloni R J, Rodriguez Batiller M J, Casal J J.2008. Abscisic acid, high-light, and oxidative stress down-regulate a photosynthetic gene via a promoter motif not involved in phytochrome-mediated transcriptional regulation[J]. Molecular Plant, 1(1): 75-83. [88] Tada Y, Sakamoto M, Matsuoka M, et al.1991. Expression of a monocot LHCP promoter in transgenic rice[J]. The EMBO Journal, 10(7): 1803-1808. [89] Timperio A M, Gevi F, Ceci L R, et al.2012. Acclimation to intense light implies changes at the level of trimeric subunits involved in the structural organization of the main light-harvesting complex of photosystem II (LHCII) and their isoforms[J]. Plant Physiology & Biochemistry, 50(1): 8-14. [90] Voigt C, Oster U, Börnke F, et al.2010. In-depth analysis of the distinctive effects of norflurazon implies that tetrapyrrole biosynthesis, organellar gene expression and ABA cooperate in the GUN-type of plastid signalling[J]. Physiologia Plantarum, 138(4): 503-519. [91] Wang J Y, Chen J D, Wang S L, et al.2020. Repressed gene expression of photosynthetic antenna proteins associated with yellow leaf variation as revealed by bulked segregant RNA-seq in tea plant Camellia sinensis[J]. Journal of Agricultural and Food Chemistry, 68(30): 8068-8079. [92] Wang S, Wang P, Gao L, et al.2017. Characterization and complementation of a chlorophyll-less dominant mutant GL1 in Lagerstroemia indica[J]. DNA and Cell Biology, 36(5): 354-366. [93] Wei Y, Lu X, Bao J, et al.2022. Identification and expression analysis of chlorophyll a/b binding protein gene family in grape (Vitis vinifera)[J]. Physiology and Molecular Biology of Plants, 28(6): 1147-1158. [94] Wientjes E, Oostergetel G T, Jansson S, et al.2009. The role of Lhca complexes in the supramolecular organization of higher plant photosystem I[J]. Journal of Biological Chemistry, 284(12): 7803-7810. [95] Wu P, Kong Q, Bian J, et al.2022. Unveiling molecular mechanisms of nitric oxide-induced low-temperature tolerance in cucumber by transcriptome profiling[J]. International Journal of Molecular Sciences, 23(10): 5615. [96] Xia Y, Ning Z, Bai G, et al.2012. Allelic variations of a light harvesting chlorophyll a/b binding protein gene (Lhcb1) associated with agronomic traits in barley[J]. The Public Library of Science One, 7(5): e37573. [97] Xu Y H, Liu R, Yan L, et al.2012. Light-harvesting chlorophyll a/b binding proteins are required for stomatal response to abscisic acid in Arabidopsis[J]. Journal of Experimental Botany, 63(3): 1095-1106. [98] Zhang M, Senoura T, Yang X, et al.2011. Lhcb2 gene expression analysis in two ecotypes of Sedum alfredii subjected to Zn/Cd treatments with functional analysis of SaLhcb2 isolated from a Zn/Cd hyperaccumulator[J]. Biotechnology Letters, 33(9): 1865-1871. [99] Zhang Q, Ma C, Wang X, et al.2021. Genome‐wide identification of the light‐harvesting chlorophyll a/b binding (Lhc) family in Gossypium hirsutum reveals the influence of GhLhcb2. 3 on chlorophyll a synthesis[J]. Plant Biology, 23(5): 831-842. [100] Zhao S, Gao H, Luo J, et al.2020. Genome-wide analysis of the light-harvesting chlorophyll a/b binding gene family in apple (Malus domestica) and functional characterization of MdLhcb4.3, which confers tolerance to drought and osmotic stress[J]. Plant Physiology and Biochemistry, 154: 517-529. [101] Zhao Y, Kong H, Guo Y, et al.2020. Light-harvesting chlorophyll a/b binding protein-coding genes in jatropha and the comparison with castor, cassava and Arabidopsis[J]. PeerJ, 8: e8465. [102] Zou Z, Li M, Jia R, et al.2020. Genes encoding light-harvesting chlorophyll a/b binding proteins in papaya (Carica papaya) and insight into lineage-specific evolution in Brassicaceae[J]. Gene, 748: 144685.