黑暗诱导衰老对不同年代冬小麦品种旗叶光系统Ⅱ功能的影响
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摘要
小麦品种的更新换代是小麦产量不断提高的重要因素,阐明小麦品种演替过程中不同生理特性的变化对新品种选育具有重要参考价值.旗叶衰老速率快慢是影响小麦产量水平的关键因素,目前对于不同小麦品种衰老过程中旗叶光系统Ⅱ功能的变化规律尚不清楚.本试验选用1941—2014年间河南地区不同时期种植的31个品种,通过黑暗诱导离体叶片衰老,测定旗叶叶绿素荧光诱导动力学参数、叶绿素相对含量的变化,分析了光系统Ⅱ功能的变化规律.结果表明:品种演替过程中旗叶的叶绿素含量逐渐提高,衰老过程中近代品种叶绿素的降解速率低于较早年代品种;旗叶衰老过程中,近代品种荧光诱导动力学曲线的J点上升幅度小于I点;品种更替过程中光系统Ⅱ最大光化学效率和单位面积有活性反应中心数目逐渐增加,但是近代品种降低速率低于较早年代品种.叶绿素含量的变化与未衰老叶片中F_v/F_m没有显著相关性,但是随着衰老程度增加,相关性逐渐增大,且趋势线斜率逐渐提高;光系统Ⅱ单位面积有活性反应中心数目与品种育成时间呈显著正相关,且随着衰老程度增加,相关程度和趋势线斜率均显著提高.综上,小麦品种演替过程中,旗叶叶绿素含量逐渐升高,降解速率逐渐减缓,光合电子传递过程中Q_A到Q_B电子传递的抗衰老能力得到改善,从而减缓了衰老过程中光系统Ⅱ最大光化学效率和有活性反应中心的衰减速率,同时,叶绿素含量的提高和旗叶光系统Ⅱ抗衰老能力的增强也是品种更替过程中产量逐渐提高的重要因素.
Cultivar renewing is important for the increases of wheat yield. Studying changes of different physiological characteristics in the succession process of wheat varieties has great implications for future breeding. The senescence rate of flag leaf is a key factor affecting winter wheat yield. The variation of photosystem Ⅱ function during senescence of flag leaves of wheat from different ages is still not clear. 31 wheat varieties planted in Henan Province from different ages since 1941 were examined in this experiment. The variation of photosystem Ⅱ function was analyzed through measuring the relative chlorophyll content,and chlorophyll fluorescence induction dynamics during the senescence of flag leaves which were induced by continued dark. The results showed that the chlorophyll content of flag leaves was gradually increased in the succession of winter wheat. The chlorophyll degradation rate in the leaves of modern varieties was lower than the earlier varieties during the senescnece of flag leaves. Meanwhile,J point of the fluorescence induction kinetics curves in flag leaves of modern varieties increased less than I point. The photosystem Ⅱ maximum photochemical efficiency and the amount of active reaction centers per unit area gradually increased during succession of wheat varieties,but the reduced extent in leaves of modern varieties was lower than that in the earlier ones. There was no significant correlation between the change of chlorophyll content and F_v/F_(m) in senescent leaves. There was significant positive correlation when the leaves aging degree increased,with the slope of trend line gradually increased. The photosystem Ⅱ unit area and the amount of active reaction center was positively related with the aging degree,and both the correlation degree and slope of trend line increased with the increases of senescence extent of the leaves. The results suggested that chlorophyll content of flag leaf increased and the degradation rate slowed down gradually during the succession of wheat varieties. The anti-aging ability of photosynthetic electron transfered from Q_A to Q_Bwas improved,which contributed to the slowing down of the decline of F_v/F_m and the amount of active reaction center. The increased chlorophyll content and anti-aging ability of photosystem Ⅱ also contributed to the increases of yield in the succession of winter wheat.
Cultivar renewing is important for the increases of wheat yield. Studying changes of different physiological characteristics in the succession process of wheat varieties has great implications for future breeding. The senescence rate of flag leaf is a key factor affecting winter wheat yield. The variation of photosystem Ⅱ function during senescence of flag leaves of wheat from different ages is still not clear. 31 wheat varieties planted in Henan Province from different ages since 1941 were examined in this experiment. The variation of photosystem Ⅱ function was analyzed through measuring the relative chlorophyll content,and chlorophyll fluorescence induction dynamics during the senescence of flag leaves which were induced by continued dark. The results showed that the chlorophyll content of flag leaves was gradually increased in the succession of winter wheat. The chlorophyll degradation rate in the leaves of modern varieties was lower than the earlier varieties during the senescnece of flag leaves. Meanwhile,J point of the fluorescence induction kinetics curves in flag leaves of modern varieties increased less than I point. The photosystem Ⅱ maximum photochemical efficiency and the amount of active reaction centers per unit area gradually increased during succession of wheat varieties,but the reduced extent in leaves of modern varieties was lower than that in the earlier ones. There was no significant correlation between the change of chlorophyll content and F_v/F_(m) in senescent leaves. There was significant positive correlation when the leaves aging degree increased,with the slope of trend line gradually increased. The photosystem Ⅱ unit area and the amount of active reaction center was positively related with the aging degree,and both the correlation degree and slope of trend line increased with the increases of senescence extent of the leaves. The results suggested that chlorophyll content of flag leaf increased and the degradation rate slowed down gradually during the succession of wheat varieties. The anti-aging ability of photosynthetic electron transfered from Q_A to Q_Bwas improved,which contributed to the slowing down of the decline of F_v/F_m and the amount of active reaction center. The increased chlorophyll content and anti-aging ability of photosystem Ⅱ also contributed to the increases of yield in the succession of winter wheat.
引文
[1]State Statistical Bureau(国家统计局).China Statistic Yearbook.Beijing:China Statistics Press,2015(in Chinese)
[2]State Statistical Bureau(国家统计局).China Statistic Yearbook.Beijing:China Statistics Press,2016(in Chinese)
[3]Zhu C-C(朱灿灿),Zhang B-N(张宝娜),Wang X(王翔),et al.Analysis of grain filling characteristics of wheat cultivars from different ages in Henan Province.Journal of Henan Agricultural Sciences(河南农业科学),2014,43(4):32-35(in Chinese)
[4]Yang W-G(杨武广),Tian Z-W(田中伟),Yin M(殷美),et al.Variations of grain yield and quality characteristics in winter wheat cultivars from different eras and their responses to nitrogen.Journal of Triticeae Crops(麦类作物学报),2014,34(10):1390-1397(in Chinese)
[5]Wang Y-Q(王义芹),Tan W(谭伟),Yang X-H(杨兴洪),et al.Photosynthetic characteristics and the activities of antioxidative enzymes in flag leaves of wheat cultivars released in different period.Acta Botanica Boreali-Occidentalia Sinica(西北植物学报),2007,27(12):2484-2490(in Chinese)
[6]Yang Z-J(杨再洁),Chen F(陈阜),Shi L-G(史磊刚),et al.Responses of photosynthetic characteristics of winter wheat cultivars released in different decades to water deficit in North China Plain.Acta Agronomica Sinica(作物学报),2013,39(4):693-703(in Chinese)
[7]Yan C-S(闫长生),Zhang H-P(张海萍),Hai L(海林),et al.Differences of preharvest sprouting resistance among Chinese wheat cultivars.Acta Agronomica Sinica(作物学报),2006,32(4):580-587(in Chinese)
[8]Meng W-W(孟维伟),Zhang Y-L(张永丽),Ma XH(马兴华),et al.Effects of irrigation stage and amount on water consumption characteristics,flag leaf photosynthesis,and grain yield in wheat.Acta Agronomica Sinica(作物学报),2009,35(10):1884-1892(in Chinese)
[9]Xu L(徐澜),Gao Z-Q(高志强),An W(安伟),et al.Flag leaf photosynthetic characteristics,change in chlorophyll fluorescence parameters,and their relationships with yield of winter wheat sowed in spring.Chinese Journal of Applied Ecology(应用生态学报),2016,27(1):133-142(in Chinese)
[10]Lee IH,Lee IC,Kim J,et al.NORE1/SAUL1integrates temperature-dependent defense programs involving SGT1b and PAD4 pathways and leaf senescence in Arabidopsis.Physiologia Plantarum,2016,158:180-199
[11]Zhang L-P(张黎萍),Jing Q(荆奇),Dai T-B(戴廷波),et al.Effects of temperature and illumination on flag leaf photosynthetic characteristics and senescence of wheat cultivars with different grain quality.Chinese Journal of Applied Ecology(应用生态学报),2008,19(2):311-316(in Chinese)
[12]Naschitz S,Naor A,Wolf S,et al.The effects of temperature and drought on autumnal senescence and leaf shed in apple under warm,east mediterranean climate.Trees,2014,28:879-890
[13]Lu Q,Lu C,Zhang J,et al.Photosynthesis and chlorophyll a fluorescence during flag leaf senescence of fieldgrown wheat plants.Journal of Plant Physiology,2002,159:1173-1178
[14]Kotakis C,Theodoropoulou E,Tassis K,et al.Putrescine,a fast-acting switch for tolerance against osmotic stress.Journal of Plant Physiology,2014,171:48-51
[15]Weaver LM,Amasion RM.Senescence is induced in individually darkened Arabidopsis leaves,but inhibited in whole darkened plants.Plant Physiology,2001,127:876-886
[16]Liebsch D,Keech O.Dark-induced leaf senescence:New insights into a complex light-dependent regulatory pathway.New Phytologist,2016,212:563-570
[17]Wada S,Ishida H,Izumi M,et al.Autophagy plays a role in chloroplast degradation during senescence in individually darkened leaves.Plant Physiology,2009,149:885-893
[18]Yang C,Zhang Z,Gao H,et al.Mechanisms by which the infection of Sclerotinia sclerotiorum(Lib.)de Bary affects the photosynthetic performance in tobacco leaves.BMC Plant Biology,2014,14:240
[19]Zhang Z-S(张子山),Zhang L-T(张立涛),Gao H-Y(高辉远),et al.Research of the photoinhibition of PSI and PSⅡin leaves of cucumber under chilling stress combined with different light intensities.Scientia Agricultura Sinica(中国农业科学),2009,42(12):4288-4293(in Chinese)
[20]Schansker G,Srivastava A,Strasser RJ.Characterization of the 820-nm transmission signal paralleling the chlorophyll a fluorescence rise(OJIP)in pea leaves.Functional Plant Biology,2003,30:785-796
[21]Li P-M(李鹏民),Gao H-Y(高辉远),Strasser RJ.Application of the fast chlorophyll fluorescence induction dynamics analysis in photosynthesis study.Journal of Plant Physiology and Molecular Biology(植物生理与分子生物学学报),2005,31(6):559-566(in Chinese)
[22]Chang SH,Ying J,Zhang JJ,et al.Expression of a wheat S-like RNnase(WRN1)c DNA during natural and dark-induced senescence.Acta Botanica Sinica,2003,45:1071-1075
[23]Buchanan-Wollaston V,Page T,Harrison E,et al.Comparative transcriptome analysis reveals significant differences in gene expression and signalling pathways between developmental and dark/starvation-induced senescence in Arabidopsis.Plant Journal,2005,42:567-585
[24]Kanazawa S,Sano S,Koshiba T,et al.Changes in antioxidative enzymes in cucumber cotyledons during natural senescence:Comparison with those during darkinduced senescence.Physiologia Plantarum,2000,109:211-216
[25]Driever SM,Lawson T,Andralojc PJ,et al.Natural variation in photosynthetic capacity,growth,and yield in 64 field-grown wheat genotypes.Journal of Experimental Botany,2014,65:4959-4973
[26]Wang S-H(王士红),Jin Q(荆奇),Dai T-B(戴廷波),et al.Evolution characteristics of flag leaf photosynthesis and grain yield of wheat cultivars bred in different years.Chinese Journal of Applied Ecology(应用生态学报),2008,19(6):1255-1260(in Chinese)
[27]Long Y-R(龙元薷),Liu J-G(刘建国),Zhang L-T(张立涛).The cell growth enhancement and difference of 4 strains of Haematococcus pluvialis under mixotrophic culture mode.Marine Sciences(海洋科学),2014,38(12):1-7(in Chinese)
[28]Fan X,Zhang Z,Gao H,et al.Photoinhibition-like damage to the photosynthetic apparatus in plant leaves induced by submergence treatment in the dark.PLo S One,2014,9(2):e89067
[29]Yang C,Zhang Z,Gao H,et al.The mechanism by which Na Cl treatment alleviates PSI photoinhibition under chilling-light treatment.Journal of Photochemistry and Photobiology B:Biology,2014,140:286-291
[30]Zhang Z-S(张子山),Yang C(杨程),Gao H-Y(高辉远),et al.Relationship between photochemistry activity and decrease in chlorophyll content during senescence in leaves of stay green and quick-leaf-senescence inbred line of maize.Scientia Agricultura Sinica(中国农业科学),2012,45(23):4794-4800(in Chinese)
[31]Gao J,Li P,Ma F,et al.Photosynthetic performance during leaf expansion in Malus micromalus probed by chlorophyll a fluorescence and modulated 820 nm reflection.Journal of Photochemistry and Photobiology B:Biology,2014,137:144-150
[32]Yang D-Q(杨东清),Wang Z-L(王振林),Yin Y-P(尹燕枰),et al.Effects of exogenous ABA and 6-BA on flag leaf senescence in different types of stay-green wheat and relevant physiological mechanisms.Acta Agronomica Sinica(作物学报),2013,39(6):1096-1104(in Chinese)
[33]Zhao C-X(赵长星),Ma D-H(马东辉),Wang Y-F(王月福),et al.Effect of nitrogen application rate and post-anthesis soil moisture content on the flag leaf senescence and kernel weight of wheat.Chinese Journal of Applied Ecology(应用生态学报),2008,19(11):2388-2393(in Chinese)
[2]State Statistical Bureau(国家统计局).China Statistic Yearbook.Beijing:China Statistics Press,2016(in Chinese)
[3]Zhu C-C(朱灿灿),Zhang B-N(张宝娜),Wang X(王翔),et al.Analysis of grain filling characteristics of wheat cultivars from different ages in Henan Province.Journal of Henan Agricultural Sciences(河南农业科学),2014,43(4):32-35(in Chinese)
[4]Yang W-G(杨武广),Tian Z-W(田中伟),Yin M(殷美),et al.Variations of grain yield and quality characteristics in winter wheat cultivars from different eras and their responses to nitrogen.Journal of Triticeae Crops(麦类作物学报),2014,34(10):1390-1397(in Chinese)
[5]Wang Y-Q(王义芹),Tan W(谭伟),Yang X-H(杨兴洪),et al.Photosynthetic characteristics and the activities of antioxidative enzymes in flag leaves of wheat cultivars released in different period.Acta Botanica Boreali-Occidentalia Sinica(西北植物学报),2007,27(12):2484-2490(in Chinese)
[6]Yang Z-J(杨再洁),Chen F(陈阜),Shi L-G(史磊刚),et al.Responses of photosynthetic characteristics of winter wheat cultivars released in different decades to water deficit in North China Plain.Acta Agronomica Sinica(作物学报),2013,39(4):693-703(in Chinese)
[7]Yan C-S(闫长生),Zhang H-P(张海萍),Hai L(海林),et al.Differences of preharvest sprouting resistance among Chinese wheat cultivars.Acta Agronomica Sinica(作物学报),2006,32(4):580-587(in Chinese)
[8]Meng W-W(孟维伟),Zhang Y-L(张永丽),Ma XH(马兴华),et al.Effects of irrigation stage and amount on water consumption characteristics,flag leaf photosynthesis,and grain yield in wheat.Acta Agronomica Sinica(作物学报),2009,35(10):1884-1892(in Chinese)
[9]Xu L(徐澜),Gao Z-Q(高志强),An W(安伟),et al.Flag leaf photosynthetic characteristics,change in chlorophyll fluorescence parameters,and their relationships with yield of winter wheat sowed in spring.Chinese Journal of Applied Ecology(应用生态学报),2016,27(1):133-142(in Chinese)
[10]Lee IH,Lee IC,Kim J,et al.NORE1/SAUL1integrates temperature-dependent defense programs involving SGT1b and PAD4 pathways and leaf senescence in Arabidopsis.Physiologia Plantarum,2016,158:180-199
[11]Zhang L-P(张黎萍),Jing Q(荆奇),Dai T-B(戴廷波),et al.Effects of temperature and illumination on flag leaf photosynthetic characteristics and senescence of wheat cultivars with different grain quality.Chinese Journal of Applied Ecology(应用生态学报),2008,19(2):311-316(in Chinese)
[12]Naschitz S,Naor A,Wolf S,et al.The effects of temperature and drought on autumnal senescence and leaf shed in apple under warm,east mediterranean climate.Trees,2014,28:879-890
[13]Lu Q,Lu C,Zhang J,et al.Photosynthesis and chlorophyll a fluorescence during flag leaf senescence of fieldgrown wheat plants.Journal of Plant Physiology,2002,159:1173-1178
[14]Kotakis C,Theodoropoulou E,Tassis K,et al.Putrescine,a fast-acting switch for tolerance against osmotic stress.Journal of Plant Physiology,2014,171:48-51
[15]Weaver LM,Amasion RM.Senescence is induced in individually darkened Arabidopsis leaves,but inhibited in whole darkened plants.Plant Physiology,2001,127:876-886
[16]Liebsch D,Keech O.Dark-induced leaf senescence:New insights into a complex light-dependent regulatory pathway.New Phytologist,2016,212:563-570
[17]Wada S,Ishida H,Izumi M,et al.Autophagy plays a role in chloroplast degradation during senescence in individually darkened leaves.Plant Physiology,2009,149:885-893
[18]Yang C,Zhang Z,Gao H,et al.Mechanisms by which the infection of Sclerotinia sclerotiorum(Lib.)de Bary affects the photosynthetic performance in tobacco leaves.BMC Plant Biology,2014,14:240
[19]Zhang Z-S(张子山),Zhang L-T(张立涛),Gao H-Y(高辉远),et al.Research of the photoinhibition of PSI and PSⅡin leaves of cucumber under chilling stress combined with different light intensities.Scientia Agricultura Sinica(中国农业科学),2009,42(12):4288-4293(in Chinese)
[20]Schansker G,Srivastava A,Strasser RJ.Characterization of the 820-nm transmission signal paralleling the chlorophyll a fluorescence rise(OJIP)in pea leaves.Functional Plant Biology,2003,30:785-796
[21]Li P-M(李鹏民),Gao H-Y(高辉远),Strasser RJ.Application of the fast chlorophyll fluorescence induction dynamics analysis in photosynthesis study.Journal of Plant Physiology and Molecular Biology(植物生理与分子生物学学报),2005,31(6):559-566(in Chinese)
[22]Chang SH,Ying J,Zhang JJ,et al.Expression of a wheat S-like RNnase(WRN1)c DNA during natural and dark-induced senescence.Acta Botanica Sinica,2003,45:1071-1075
[23]Buchanan-Wollaston V,Page T,Harrison E,et al.Comparative transcriptome analysis reveals significant differences in gene expression and signalling pathways between developmental and dark/starvation-induced senescence in Arabidopsis.Plant Journal,2005,42:567-585
[24]Kanazawa S,Sano S,Koshiba T,et al.Changes in antioxidative enzymes in cucumber cotyledons during natural senescence:Comparison with those during darkinduced senescence.Physiologia Plantarum,2000,109:211-216
[25]Driever SM,Lawson T,Andralojc PJ,et al.Natural variation in photosynthetic capacity,growth,and yield in 64 field-grown wheat genotypes.Journal of Experimental Botany,2014,65:4959-4973
[26]Wang S-H(王士红),Jin Q(荆奇),Dai T-B(戴廷波),et al.Evolution characteristics of flag leaf photosynthesis and grain yield of wheat cultivars bred in different years.Chinese Journal of Applied Ecology(应用生态学报),2008,19(6):1255-1260(in Chinese)
[27]Long Y-R(龙元薷),Liu J-G(刘建国),Zhang L-T(张立涛).The cell growth enhancement and difference of 4 strains of Haematococcus pluvialis under mixotrophic culture mode.Marine Sciences(海洋科学),2014,38(12):1-7(in Chinese)
[28]Fan X,Zhang Z,Gao H,et al.Photoinhibition-like damage to the photosynthetic apparatus in plant leaves induced by submergence treatment in the dark.PLo S One,2014,9(2):e89067
[29]Yang C,Zhang Z,Gao H,et al.The mechanism by which Na Cl treatment alleviates PSI photoinhibition under chilling-light treatment.Journal of Photochemistry and Photobiology B:Biology,2014,140:286-291
[30]Zhang Z-S(张子山),Yang C(杨程),Gao H-Y(高辉远),et al.Relationship between photochemistry activity and decrease in chlorophyll content during senescence in leaves of stay green and quick-leaf-senescence inbred line of maize.Scientia Agricultura Sinica(中国农业科学),2012,45(23):4794-4800(in Chinese)
[31]Gao J,Li P,Ma F,et al.Photosynthetic performance during leaf expansion in Malus micromalus probed by chlorophyll a fluorescence and modulated 820 nm reflection.Journal of Photochemistry and Photobiology B:Biology,2014,137:144-150
[32]Yang D-Q(杨东清),Wang Z-L(王振林),Yin Y-P(尹燕枰),et al.Effects of exogenous ABA and 6-BA on flag leaf senescence in different types of stay-green wheat and relevant physiological mechanisms.Acta Agronomica Sinica(作物学报),2013,39(6):1096-1104(in Chinese)
[33]Zhao C-X(赵长星),Ma D-H(马东辉),Wang Y-F(王月福),et al.Effect of nitrogen application rate and post-anthesis soil moisture content on the flag leaf senescence and kernel weight of wheat.Chinese Journal of Applied Ecology(应用生态学报),2008,19(11):2388-2393(in Chinese)