外源6-BA对紫花苜蓿盛花期叶片光合、生理特性及结荚率的影响
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摘要
为了探讨外源6-苄氨基嘌呤(6-BA)对紫花苜蓿(Medicago sativa)叶片光合作用、抗氧化系统及结荚率的调控作用,以新牧4号紫花苜蓿(Medicago sativa ‘Xinmu No.4’)为试验材料,现蕾期叶面喷施10、20、30 mg·L~(-1)的6-BA,以喷等量清水为对照,研究外源6-BA对苜蓿盛花期叶片光合参数、荧光参数、抗氧化酶活性、渗透调节物质及结荚率的影响。结果表明,20 mg·L~(-1)的6-BA处理显著(P<0.05)提高了苜蓿叶片的净光合速率(P_n)、蒸腾速率(T_r)、最大光化学效率(F_v/F_m),增强了苜蓿叶片中超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)的活性,使叶片中积累了脯氨酸和可溶性糖,减少了叶片中过氧化氢的积累,显著提高了苜蓿的结荚率及单株种子产量(P<0.05);~130 mg·L–处理,虽提高苜蓿叶片抗氧化酶活性,并且主要荧光参数高于对照,但Pn低于对照,该处理中P_n的下降由非气孔限制因素主导。20 mg·L~(-1)的6-BA可提高盛花期苜蓿叶片的光合速率,抗氧化酶活性,减小光合机构的损伤,有利于苜蓿叶片光合作用的进行,提高苜蓿的结荚率及种子产量。
The role of exogenous 6-benzylaminopurine (6-BA) in the regulation of photosynthesis and the antioxidant system in alfalfa leaves was evaluated to provide a theoretical and research basis the application of this cytokinin in the growth and development of alfalfa. Xinmu No. 4 alfalfa(Medicago sativa ‘Xinmu No. 4') plants were examined in this study. 6-BA solutions at concentrations of 10, 20, and 30 mg·L~(-1) were sprayed onto the leaves of alfalfa during the squaring period; the control plants were sprayed with water. The effects of exogenous 6-BA on photosynthetic parameters, fluorescence parameters, antioxidant enzyme activities, osmotic adjustment substances, and rate of podding were examined during the flowering stage of alfalfa. The results showed that after treatment with 20 mg·L~(-1) 6-BA, the photosynthetic rate, transpiration rate, and maximum photochemical efficiency of the alfalfa leaves were significantly increased and the activities of superoxide dismutase, peroxidase, and catalase in the leaves of alfalfa were enhanced. The increase accumulation of proline and soluble sugars and the reduced accumulation of hydrogen peroxide in the leaves significantly improved the fruiting rate and seed yield per plant. In the 30 mg·L~(-1) treatment, although the antioxidative enzyme activity was increased and the main fluorescence parameters were higher than those of the control, the rate of photosynthesis was decreased compared to the control. The decrease in the photosynthetic rate under this treatment was dominated by non-stomatal limiting factors. In conclusion, 20 mg·L~(-1) 6-BA can increase the photosynthetic rate and antioxidant enzyme activity as well as reduce damage to the photosynthetic mechanism of alfalfa leaves at the flowering stage. Moreover, increasing photosynthesis in the alfalfa leaves can increase the fruiting rate and seed yield.
The role of exogenous 6-benzylaminopurine (6-BA) in the regulation of photosynthesis and the antioxidant system in alfalfa leaves was evaluated to provide a theoretical and research basis the application of this cytokinin in the growth and development of alfalfa. Xinmu No. 4 alfalfa(Medicago sativa ‘Xinmu No. 4') plants were examined in this study. 6-BA solutions at concentrations of 10, 20, and 30 mg·L~(-1) were sprayed onto the leaves of alfalfa during the squaring period; the control plants were sprayed with water. The effects of exogenous 6-BA on photosynthetic parameters, fluorescence parameters, antioxidant enzyme activities, osmotic adjustment substances, and rate of podding were examined during the flowering stage of alfalfa. The results showed that after treatment with 20 mg·L~(-1) 6-BA, the photosynthetic rate, transpiration rate, and maximum photochemical efficiency of the alfalfa leaves were significantly increased and the activities of superoxide dismutase, peroxidase, and catalase in the leaves of alfalfa were enhanced. The increase accumulation of proline and soluble sugars and the reduced accumulation of hydrogen peroxide in the leaves significantly improved the fruiting rate and seed yield per plant. In the 30 mg·L~(-1) treatment, although the antioxidative enzyme activity was increased and the main fluorescence parameters were higher than those of the control, the rate of photosynthesis was decreased compared to the control. The decrease in the photosynthetic rate under this treatment was dominated by non-stomatal limiting factors. In conclusion, 20 mg·L~(-1) 6-BA can increase the photosynthetic rate and antioxidant enzyme activity as well as reduce damage to the photosynthetic mechanism of alfalfa leaves at the flowering stage. Moreover, increasing photosynthesis in the alfalfa leaves can increase the fruiting rate and seed yield.
引文
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[2]GOMEZ-LOBATO M E,CIVELLO P M,MARTíNEZ G A.Effects of ethylene,cytokinin and physical treatments on BoPaO gene expression of harvested broccoli.Journal of the Science of Food&Agriculture,2012,92(1):151-158.
[3]RUSHING J W.Cytokinins affect respiration,ethylene production,and chlorophyll retention of packaged broccoli florets.Hortscience A Publication of the American Society for Horticultural Science,1990,25(1):88-90.
[4]XU F,YANG Z,CHEN X,JIN P,WANG X,ZHEN Y.6-Benzylaminopurine delays senescence and enhances health-promoting compounds of harvested broccoli.Journal of Agricultural&Food Chemistry,2012,60(1):234-240.
[5]徐运清,李顺文,鲁旭东.6-BA对小麦叶片几种保护酶活性的影响.农业与技术,2000,20(4):13-15.XU Y Q,LI S W,LU X D.Effect of 6-BA on several protective enzyme activity of wheat leaf.Agriculture&Technology,2000,20(4):13-15.
[6]张治安,陈庆利,徐克章.灌浆期喷施6-BA对甜玉米一些生理指标的影响.植物生理学通讯,2004,40(6):683-685.ZHANG Z A,CHEN Q L,XU K Z.Effects of foliage spray with 6-BA during filling stage on some physiologlcal indexes of sugar maize.Plant Physiology Journal,2004,40(6):683-685.
[7]董永华,史吉平,周慧欣.6-BA对小麦幼苗抗旱性的影响.植物营养与肥料学报,1999,5(1):73-76.DONG Y H,SHI J P,ZHOU H X.Effect of 6-BA on drought resistance in wheat seedlings.Plant Nutrition and Fertilizer Science,1999,5(1):73-76.
[8]SHAO R,WANG K,SHANGGUAN Z.Cytokinin-induced photosynthetic adaptability of Zea mays L.to drought stress associated with nitric oxide signal:Probed by ESR spectroscopy and fast OJIP fluorescence rise.Journal of Plant Physiology,2010,167(6):472-475.
[9]WU X,ZHU Z,LI X,ZHA D.Effects of cytokinin on photosynthetic gas exchange,chlorophyll fluorescence parameters and antioxidative system in seedlings of eggplant(Solanum melongena L.)under salinity stress.Acta Physiologiae Plantarum,2012,34(6):2105-2114.
[10]杨芳芳,张国斌,颉建明,郁继华.6-BA预处理对低温弱光胁迫下辣椒幼苗叶绿素a荧光参数和膜脂过氧化的影响.植物生理学通讯,2009,45(6):575-578.YANG F F,ZHANG G B,JIE J M,YU J H.Effects of 6-BA pretreatment on chlorophyll fluorescence parameter and lipid peroxodatio in pepperseedlings under low temperature and weak light stress.Plant Physiology Journal,2009,45(6):575-578.
[11]NAGEL L,BREWSTER R,RIEDELL W E,REESE R N.Cytokinin Regulation of Flower and Pod Set in Soybeans(Glycine max L.Merr.).Annals of Botany,2001,88(1):27-31.
[12]ATKINS C A,PIGEAIRE A.Application of cytokinins to flowers to increase pod set in Lupinus angustifolius.Australian Journal of Agricultural Research,1993,44:1799-1819.
[13]PAN S,Fahd R,Li W,TIAN H,MO Z,DUAN M.Roles of plant growth regulators on yield,grain qualities and antioxidant enzyme activities in super hybrid rice(Oryza sativa L.).Rice,2013,6(1):9-11.
[14]魏小星,郭文山,孙彦.生长调节剂对紫花苜蓿种子产量及产量构成要素的影响.草业科学,2009,26(6):121-125.WEI X X,GUO W S,SUN Y.Effects of growth regulators on seed yield and yield components of alfalfa.Pratacultural Science,2009,26(6):121-125.
[15]周万福,张亚玲,刘香萍.植物生长调节剂对紫花苜蓿种子产量及其构成因子的影响.当代畜牧,2011(4):45-47.ZHOU W F,ZHANG Y L,LIU X P.Effects of plant growth regulator on seed yield and component factors of alfalfa.Contemporary Animal Husbandry,2011(4):45-47.
[16]王玉祥.激素与水肥对紫花苜蓿生殖生长及种子产量的影响.乌鲁木齐:新疆农业大学博士学位论文,2015.WANG Y X.Effects of hormones and water fertilizer on reproductive growth and seed yield of alfalfa.PhD Thesis.Urumqi:Xinjiang Agricultural University,2015.
[17]张志良.植物生理学实验指导(第二版).北京:高等教育出版社,1990:45-67.ZHANG Z L.Experimental Instruction in Plant Physiology(second edition).Beijing:Advanced Education Press,1990:45-67.
[18]张海娜,李存东,肖凯.外源6-BA对棉花光合和叶片衰老特性的调控效应研究.棉花学报,2007,19(6):467-471.ZHANG H N,LI C D,XIAO K.Regulation effects of exogenous 6-BA on photosynthesis and leaf senescence in cotton.Cotton Science,2007,19(6):467-471.
[19]王海波,王帅,王孝娣,史祥宾,王宝亮,郑晓翠,王志强,冀晓昊,刘凤之.6-BA及氨基酸硒对葡萄叶片衰老的影响.果树学报,2017,34(11):1426-1434.WANG H B,WANG S,WANG X D,SHI X B,WANG B L,ZHENG X C,WANG Z Q,JI X H.Effects of 6-BA and amino acid selenium on leaf senescence and endogenous hormones content in grape.Journal of Fruit Science,2017,34(11):1426-1434.
[20]商振清,李爱丽,董永华.6-BA或KT对水分胁迫条件下小麦旗叶光合能力及千粒重的影响.河北农业大学学报,2000,23(2):20-24.SHANG Z Q,LI A L,DONG Y H.Effect of 6-BA or KT on photosynthetic capacity in wheat flag leaf and the weight of 1000grains under water stress.Journal of Agricultural University of Hebei,2000,23(2):20-24.
[21]吴雪霞,杨晓春,朱宗文,查丁石,许爽.外源6-BA对低温胁迫下茄子幼苗光合作用、叶绿素荧光参数及光能分配的影响.植物生理学报,2013,49(11):1181-1188.WU X X,YANG X C,ZHU Z W,CHA D S,XU S.Effects of exogenous 6-BA on photosynthesis,chlorophyll fluorescence characteristics and the allocation of absorbed light in eggplant seedlings under low temperature stress.Plant Physiology Journal,2013,49(11):1181-1188.
[22]江力,孔小卫,张荣铣.6-苄基腺嘌呤和脱落酸对烟草光合功能衰退的影响.南京农业大学学报,2006,29(4):127-130.JIANG L,KONG X W,ZHANG R X.Effects of 6-benzyladenine and abscisic acid on the photosynthetic function decline in tobacco.Journal of Nanjing Agricultural University,2006,29(4):127-130.
[23]SAWAN Z M,MOHAMED A A,SAKR R A,TARRAD A M.Effect of kinetin concentration and methods of application on seed germination,yield components,yield and fiber properties of the Egyptian cotton(Gossypium barbadense).Environmental&Experimental Botany,2000,44(1):59-68.
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