外源SA与OPDA对花叶唐竹条纹叶片抗氧化系统的影响

doi:10.16420/j.issn.0513-353x.2025-0135

摘要/Abstract

摘要：

为解析外源水杨酸（SA）与12–氧代植物二烯酸（OPDA）在调控彩叶植物抗氧化机制中的作用，以花叶唐竹（Sinobambusa tootsik f. albostriata）不同表型的叶片为材料，进行不同浓度SA、OPDA和SA + OPDA处理，并测定丙二醛（MDA）和超氧阴离子（$\mathrm{O}_2^{\bar{.}}$）含量，超氧化物歧化酶（SOD）、过氧化物酶（POD）、过氧化氢酶（CAT）以及抗坏血酸—谷胱甘肽（AsA–GSH）循环相关抗氧化酶活性。结果表明，0.5 mmol · L^-1 SA、30 μmol · L^-1 OPDA及0.5 mmol · L^-1 SA + 30 μmol · L^-1 OPDA处理下，各表型叶片SOD、POD、CAT 活性和AsA–GSH循环酶活性显著提高，叶片谷胱甘肽（GSH）含量显著增加，有效清除叶片积累的活性氧（ROS）。进一步比较不同表型可知，条纹叶片中GSH含量显著高于全白叶和全绿叶，且条纹叶白色部分抗氧化特性明显优于绿色部分。这表明条纹叶片在色彩分区形成过程中具有差异化的氧化还原稳态调控能力，其中白色组织可能通过提升非酶抗氧化水平来弥补叶绿体缺失带来的氧化压力，从而有助于维持条纹结构的稳定性。综合分析表明，SA主要通过快速提升抗氧化酶活性和强化AsA–GSH循环，从而提高细胞整体的基础抗氧化能力；而OPDA作为脂氧素类信号，更倾向于调控膜脂过氧化及应激响应途径。二者在抗氧化途径中具有互补性，共同作用能更有效降低ROS水平，从而有助于维持条纹叶片的表型稳定。

关键词: 花叶唐竹, 叶色, 激素信号交流, 抗氧化能力

Abstract:

To elucidate the roles of exogenous salicylic acid（SA）and 12-oxo-phytodienoic acid （OPDA）in regulating the antioxidant mechanisms of variegated plants，leaves of different phenotypes of Sinobambusa tootsik f. albostriata were treated with various concentrations of SA，OPDA，and SA + OPDA. The contents of malondialdehyde（MDA），superoxide anion（$\mathrm{O}_2^{\bar{.}}$），as well as the activities of antioxidant enzymes including superoxide dismutase（SOD），peroxidase（POD），catalase（CAT），and antioxidant enzymes involved in the ascorbate-glutathione（AsA-GSH）cycle，were determined. The results showed that treatments with 0.5 mmol · L^-1 SA，30 μmol · L^-1 OPDA，or 0.5 mmol · L^-1 SA + 30 μmol · L^-1 OPDA significantly enhanced the activities of SOD，POD，CAT，and AsA–GSH cycle enzymes in all leaf phenotypes，markedly increased glutathione（GSH）levels，and effectively scavenged the accumulated reactive oxygen species（ROS）. Comparisons among phenotypes further revealed that GSH content in variegated leaves was significantly higher than in fully white or fully green leaves，and that the white sectors of variegated leaves exhibited stronger antioxidant characteristics than the green sectors. This suggests that variegated leaves possess differentiated redox homeostasis during the formation of color sectors，and that the white tissues may elevate non-enzymatic antioxidant capacity to compensate for the oxidative pressure caused by the absence of chloroplasts，thereby contributing to the stability of the variegated pattern. Overall，SA primarily enhances basal cellular antioxidant capacity by rapidly activating antioxidant enzymes and reinforcing the AsA–GSH cycle，whereas OPDA，as an oxylipin signal，tends to modulate lipid peroxidation and stress-responsive pathways. The two signals act in a complementary manner，and their combined application more effectively reduces ROS accumulation，thereby promoting the phenotypic stability of variegated leaves.

Key words: Sinobambusa tootsik f. albostriata, leaf color, hormone crosstalk, antioxidant capacity

唐瑜, 曾美音, 朱芃铠, 何天友, 郑郁善, 陈凌艳. 外源SA与OPDA对花叶唐竹条纹叶片抗氧化系统的影响[J]. 园艺学报, 2026, 53(5): 1477-1490.

TANG Yu, ZENG Meiyin, ZHU Pengkai, HE Tianyou, ZHENG Yushan, CHEN Lingyan. Effects of SA-OPDA Crosstalk on the Antioxidant System of Striped Leaves in Sinobambusa tootsik f. albostriata[J]. Acta Horticulturae Sinica, 2026, 53(5): 1477-1490.

图/表 10

图1 花叶唐竹的3种叶片表型

Fig. 1 Three phenotypes of leaves of Sinobambusa tootsik f. albostriata

图 2 SA与OPDA对花叶唐竹不同表型叶片丙二醛含量的影响 MOCK0：无处理对照，用于反映叶片在无任何处理条件下的基础氧化水平；MOCK1：培养液处理对照，其与MOCK0的差异表明培养液处理本身已对叶片产生一定程度的氧化胁迫；T1：0.5 mmol · L-1 SA；T2：2 mmol · L-1 SA；T3：30 μmol · L-1 OPDA；T4：50 μmol · L-1 OPDA；T5：0.5 mmol · L-1 SA + 30 μmol · L-1 OPDA；T6：2 mmol · L-1 SA + 50 μmol · L-1 OPDA。不同小写字母表示处理间差异显著（P < 0.05）。下同

Fig. 2 Effects of SA and OPDA on malondialdehyde（MDA）content in different phenotypes of Sinobambusa tootsik f. albostriata leaves MOCK0：Untreated control，reflecting the basal oxidative level of leaves without any treatment；MOCK1：Culture solution-treated control，the difference between MOCK1 and MOCK0 indicates that the culture solution itself imposed a certain degree of oxidative stress on the leaves. T1：0.5 mmol · L-1 SA；T2：2 mmol · L-1 SA；T3：30 μmol · L-1 OPDA；T4：50 μmol · L-1 OPDA；T5：0.5 mmol · L-1 SA + 30 μmol · L-1 OPDA；T6：2 mmol · L-1 SA + 50 μmol · L-1 OPDA. Different lowercase letters indicate significant differences among treatments（P < 0.05）. The same below

图3 SA与OPDA对花叶唐竹不同表型叶片超氧阴离子的影响

Fig. 3 Effects of SA and OPDA on superoxide anion levels in different phenotypes of Sinobambusa tootsik f. albostriata leaves

图 4 SA与OPDA对花叶唐竹不同表型叶片还原型谷胱甘肽含量的影响

Fig. 4 Effects of SA and OPDA on reduced glutathione content in different phenotypes of Sinobambusa tootsik f. albostriata leaves

图5 SA与OPDA对花叶唐竹不同表型叶片氧化型谷胱甘肽含量的影响

Fig. 5 Effects of SA and OPDA on oxidized glutathione content in different phenotypes of Sinobambusa tootsik f. albostriata leaves

图 6 SA与OPDA对花叶唐竹不同表型叶片抗坏血酸过氧化物酶活性的影响

Fig. 6 Effects of SA and OPDA on ascorbate peroxidase activity in different phenotypes of Sinobambusa tootsik f. albostriata leaves

图 7 SA与OPDA对花叶唐竹不同表型叶片谷胱甘肽还原酶活性的影响

Fig. 7 Effects of SA and OPDA on glutathione reductase activity in different phenotypes of Sinobambusa tootsik f. albostriata leaves

图 8 SA与OPDA对花叶唐竹不同表型叶片超氧化物歧化酶活性的影响

Fig. 8 Effects of SA and OPDA on superoxide dismutase activity in different phenotypes of Sinobambusa tootsik f. albostriata leaves

图 9 SA与OPDA对花叶唐竹不同表型叶片过氧化氢酶活性的影响

Fig. 9 Effects of SA and OPDA on catalase activity in different phenotypes of Sinobambusa tootsik f. albostriata leaves

图10 SA与OPDA对花叶唐竹不同表型叶片过氧化物酶活性的影响

Fig. 10 Effects of SA and OPDA on peroxidase activity in different phenotypes of Sinobambusa tootsik f. albostriata leaves

参考文献 35

[1]	Aluru M R, Zola J, Foudree A, Rodermel S R. 2009. Chloroplast photooxidation-induced transcriptome reprogramming in Arabidopsis immutans white leaf sectors. Plant Physiology, 150 (2):904-923. doi: 10.1104/pp.109.135780 URL
[2]	Ananieva E A, Christov K N, Popova L P. 2004. Exogenous treatment with salicylic acid leads to increased antioxidant capacity in leaves of barley plants exposed to paraquat. Journal of Plant Physiology, 161 (3):319-328. pmid: 15077630
[3]	Celi G, Gratao P L, Lanza M, Reis A. 2023. Physiological and biochemical roles of ascorbic acid on mitigation of abiotic stresses in plants. Plant Physiology and Biochemistry, 202:107970. doi: 10.1016/j.plaphy.2023.107970 URL
[4]	Chen L Y, Tang Y, Tao Y, Seerat A, Zhu P K, He T Y, Zheng Y S. 2024. Accumulation of salicylic acid and 12-oxo-phytodienoic acid acting on the antioxidant pathway to keep stability of striped leaves of variegated temple bamboo. Journal of the American Society for Horticultural Science, 149 (1):27-36. doi: 10.21273/JASHS05348-23 URL
[5]	Choi H, Yi T, Ha S H. 2021. Diversity of plastid types and their interconversions. Frontiers in Plant Science, 12:692024. doi: 10.3389/fpls.2021.692024 URL
[6]	Hasanuzzaman M, Bhuyan M, Anee T I, Parvin K, Nahar K, Mahmud J A, Fujita M. 2019. Regulation of ascorbate-glutathione pathway in mitigating oxidative damage in plants under abiotic stress. Antioxidants(Basel,Switzerland), 8 (9):384.
[7]	Herrera-Vasquez A, Salinas P, Holuigue L. 2015. Salicylic acid and reactive oxygen species interplay in the transcriptional control of defense genes expression. Frontiers in Plant Science, 6:171. doi: 10.3389/fpls.2015.00171 pmid: 25852720
[8]	Jimenez A G, Thirumalaikumar V P, Jander G, Fernie A R, Skirycz A. 2022. OPDA,more than just a jasmonate precursor. Phytochemistry,204:113432.
[9]	Liu W, Park S W. 2021. 12-Oxo-phytodienoic acid:a fuse and/or switch of plant growth and defense responses? Frontiers in Plant Science, 12:724079. doi: 10.3389/fpls.2021.724079 URL
[10]	Maynard D, Groger H, Dierks T, Dietz K J. 2018. The function of the oxylipin 12-oxophytodienoic acid in cell signaling,stress acclimation,and development. Journal of Experimental Botany, 69 (22):5341-5354. doi: 10.1093/jxb/ery316 pmid: 30169821
[11]	Milic D, Pantelic A, Banovic D B, Samardzic J, Vidovic M. 2023. Contrasting metabolisms in green and white leaf sectors of variegated Pelargonium zonale an integrative transcriptomic and metabolomic study. International Journal of Molecular Sciences, 24 (6):5288. doi: 10.3390/ijms24065288 URL
[12]	Miura E, Kato Y, Sakamoto W. 2010. Comparative transcriptome analysis of green/white variegated sectors in Arabidopsis yellow variegated2:responses to oxidative and other stresses in white sectors. Journal of Experimental Botany, 61 (9):2433-2445. doi: 10.1093/jxb/erq075 pmid: 20400527
[13]	Mur L A, Kenton P, Atzorn R, Miersch O, Wasternack C. 2006. The outcomes of concentration-specific interactions between salicylate and jasmonate signaling include synergy,antagonism,and oxidative stress leading to cell death. Plant Physiology, 140 (1):249-262. doi: 10.1104/pp.105.072348 URL
[14]	Nadarajah K K. 2020. ROS homeostasis in abiotic stress tolerance in plants. International Journal of Molecular Sciences, 21 (15):5208. doi: 10.3390/ijms21155208 URL
[15]	Noctor G, Reichheld J P, Foyer C H. 2018. ROS-related redox regulation and signaling in plants. Seminars in Cell & Developmental Biology,80:3-12.
[16]	Park S W, Li W, Viehhauser A, He B, Kim S, Nilsson A K, Andersson M X, Kittle J D, Ambavaram M M, Luan S, Esker A R, Tholl D, Cimini D, Ellerstrom M, Coaker G, Mitchell T K, Pereira A, Dietz K J, Lawrence C B. 2013. Cyclophilin 20-3 relays a 12-oxo-phytodienoic acid signal during stress responsive regulation of cellular redox homeostasis. Proceedings of the National Academy of Sciences of the United States of America, 110 (23):9559-9564.
[17]	Qi Xiaoqing, Gao Chengyue, Li Zhezhe, Yang Xiulian, Wang Lianggui, Yue Yuanzheng. 2025. Research progress on leaf variegation chimeras of ornamental plants. Acta Horticulturae Sinica, 52 (12):3401-3411. (in Chinese) doi: 10.16420/j.issn.0513-353x.2024-1060
	齐晓庆, 高澄玥, 李浙浙, 杨秀莲, 王良桂, 岳远征. 2025. 观赏植物叶色嵌合体的研究进展. 园艺学报, 52 (12):3401-3411. doi: 10.16420/j.issn.0513-353x.2024-1060
[18]	Qiao Q, Wu C, Cheng T, Yan Y, Zhang L, Wan Y, Wang J, Liu Q, Feng Z, Liu Y. 2022. Comparative analysis of the metabolome and transcriptome between the green and yellow-green regions of variegated leaves in a mutant variety of the tree species Pteroceltis tatarinowii. International Journal of Molecular Sciences, 23 (9):4950. doi: 10.3390/ijms23094950 URL
[19]	Raza A, Charagh S, Zahid Z, Mubarik M S, Javed R, Siddiqui M H, Hasanuzzaman M. 2021. Jasmonic acid:a key frontier in conferring abiotic stress tolerance in plants. Plant Cell Reports, 40 (8):1513-1541. doi: 10.1007/s00299-020-02614-z pmid: 33034676
[20]	Saleem M, Fariduddin Q, Castroverde C. 2021. Salicylic acid:a key regulator of redox signalling and plant immunity. Plant Physiology and Biochemistry,168:381-397.
[21]	Scandalios J G. 2002. The rise of ROS. Trends in Biochemical Sciences, 27 (9):483-486. doi: 10.1016/s0968-0004(02)02170-9 pmid: 12217524
[22]	Sun Y H, Hung C Y, Qiu J, Chen J, Kittur F S, Oldham C E, Henny R J, Burkey K O, Fan L, Xie J. 2017. Accumulation of high OPDA level correlates with reduced ROS and elevated GSH benefiting white cell survival in variegated leaves. Scientific Reports, 7:44158. doi: 10.1038/srep44158
[23]	Varsani S, Grover S, Zhou S, Koch K G, Huang P C, Kolomiets M V, Williams W P, Heng-Moss T, Sarath G, Luthe D S, Jander G, Louis J. 2019. 12-Oxo-phytodienoic acid acts as a regulator of maize defense against corn leaf aphid. Plant Physiology, 179 (4):1402-1415. doi: 10.1104/pp.18.01472 pmid: 30643012
[24]	Vidovic M, Morina F, Milic-Komic S, Vuleta A, Zechmann B, Prokic L, Veljovic J S. 2016a. Characterisation of antioxidants in photosynthetic and non-photosynthetic leaf tissues of variegated Pelargonium zonale plants. Plant Biology(Stuttgart,Germany), 18 (4):669-680.
[25]	Vidovic M, Morina F, Prokic L, Milic-Komic S, Zivanovic B, Jovanovic S V. 2016b. Antioxidative response in variegated Pelargonium zonale leaves and generation of extracellular H₂O₂ in(peri)vascular tissue induced by sunlight and paraquat. Journal of Plant Physiology,206:25-39.
[26]	Wang Aoqi. 2016. The research on the application of ornamental bamboo in the landscape of Changsha city[M. D. Dissertation]. Changsha: Central South University of Forestry and Technology. (in Chinese)
	王奥奇. 2016. 观赏竹在长沙市园林绿化中的应用研究[硕士论文]. 长沙: 中南林业科技大学.
[27]	Wang Peiyun. 2023. Effect of exogenous salicylic acid on the physiological and biochemical properties of Zanthoxylum armatum under waterlogging stress[M. D. Dissertation]. Chengdu: Sichuan Agricultural University. (in Chinese)
	王佩云. 2023. 外源水杨酸对水淹胁迫下竹叶花椒生理生化特性的影响[硕士论文]. 成都: 四川农业大学.
[28]	Wang X, Liu H, Yu F, Hu B, Jia Y, Sha H, Zhao H. 2019. Differential activity of the antioxidant defence system and alterations in the accumulation of osmolyte and reactive oxygen species under drought stress and recovery in rice(Oryza sativa L.)tillering. Scientific Reports, 9 (1):8543. doi: 10.1038/s41598-019-44958-x
[29]	Wang Yuhang, Li Dou, Wang Chunheng, Jin Xin, Chen Yajuan, Dai Zibo, Feng Lidan, Yang Jiangshan. 2024. The effect of melatonin on the subcellular reactive oxygen species metabolism during development and senescence in grape leaves. Acta Horticulturae Sinica, 51 (1):103-120. (in Chinese) doi: 10.16420/j.issn.0513-353x.2022-1214
	王宇航, 李斗, 王春恒, 金鑫, 陈亚娟, 戴子博, 冯丽丹, 杨江山. 2024. 褪黑素对葡萄叶片发育衰老过程中亚细胞活性氧代谢的影响. 园艺学报, 51 (1):103-120. doi: 10.16420/j.issn.0513-353x.2022-1214
[30]	Waszczak C, Carmody M, Kangasjarvi J. 2018. Reactive oxygen species in plant signaling. Annual Review of Plant Biology,69:209-236.
[31]	Wasternack C, Strnad M. 2016. Jasmonate signaling in plant stress responses and development-active and inactive compounds. New Biotechnology, 33 (5):604-613. doi: 10.1016/j.nbt.2015.11.001 URL
[32]	Xu Y Z, Arrieta-Montiel M P, Virdi K S, de Paula W B, Widhalm J R, Basset G J, Davila J I, Elthon T E, Elowsky C G, Sato S J, Clemente T E, Mackenzie S A. 2011. MutS HOMOLOG1 is a nucleoid protein that alters mitochondrial and plastid properties and plant response to high light. Plant Cell, 23 (9):3428-3441. doi: 10.1105/tpc.111.089136 URL
[33]	Yang W, Kang J, Liu Y, Guo M, Chen G. 2022. Effect of salicylic acid treatment on antioxidant capacity and endogenous hormones in winter jujube during shelf life. Food Chemistry, 397 (15):133788. doi: 10.1016/j.foodchem.2022.133788 URL
[34]	Yu Sanmiao. 2018. Study on the differences of photosynthesisand peroxidation system in the green and white tissues of Ananas comosus var. bracteatus[M. D. Dissertation]. Chengdu: Sichuan Agricultural University. (in Chinese)
	余三淼. 2018. 红苞凤梨嵌合叶片绿白组织光合作用及过氧化系统差异研究[硕士论文]. 成都: 四川农业大学.
[35]	Zhou Xuzixin, Yang Wei, Mao Meiqin, Xue Yanbin, Ma Jun. 2022. Identification of pigment components and key genes in carotenoid pathway in mutants of chimeric Ananas comosus var. bracteatus. Acta Horticulturae Sinica, 49 (5):1081-1091. (in Chinese) doi: 10.16420/j.issn.0513-353x.2021-0355
	周徐子鑫, 杨威, 毛美琴, 薛彦斌, 马均. 2022. 金边红苞凤梨叶色突变体色素鉴定及类胡萝卜素合成限速基因筛选. 园艺学报, 49 (5):1081-1091. doi: 10.16420/j.issn.0513-353x.2021-0355