春季低温对小麦产量和光合特性的影响
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摘要
黄淮麦区频发春季低温冻害,造成小麦减产。为探讨小麦品种对春季低温胁迫响应的生理特点,确定品种抗寒性鉴定的生理指标,以泰山6426、泰山4033和济麦22为试验材料,研究了起身期、拔节期和孕穗期低温处理下不同品种的产量性状、光合生理指标及叶片细胞结构差异。3个品种受低温胁迫后,观测农艺性状的变化趋势相似,且年度间有一定的一致性。不同低温处理对株高的影响相对较小,尤其是起身期低温处理;不同生育期低温处理,小麦单株穗数均显著低于对照,但下降幅度在年度间不完全一致;单株穗粒数和单株产量随低温处理的推迟呈现显著降低趋势,尤以孕穗期低温处理的降低幅度最大,单株穗粒数比对照低90.5%~93.3%(2012—2013)和91.9%~96.6%(2013—2014),单株产量比对照低87.9%~97.3%(2012—2013)和91.5%~97.8%(2013—2014);拔节期是低温伤害的第二敏感期,而起身期低温对小麦产量的影响相对最小。减产的主要原因是穗粒数降低。无论是拔节期还是孕穗期低温处理,3个品种的叶绿素含量均呈现下降趋势,下降的幅度因品种而异;低温胁迫下光合速率、蒸腾速率和叶片气孔导度均显著下降,胞间CO_2浓度均有较大幅度的升高,3个品种中泰山6426受影响最小。低温胁迫对叶绿素荧光参数也有明显影响,除泰山6426外,其余2个品种的F_v/F_m均显著下降。各品种受低温伤害后表皮叶肉细胞破坏严重,排列趋于松散,保卫细胞和副卫细胞变形,气孔变大。本研究结果显示,光合速率、蒸腾速率、叶片气孔导度、胞间CO_2浓度及F_v/F_m可作为评价品种春季抗寒性的生理指标。
Low temperature(LT) stress in spring occurs frequently in the Huang-Huai Rivers Valley, resulting in yield loss of wheat. The objectives of this study were to understand the physiological responses of wheat cultivars to LT and determine suitable physiological indicator(s) for cold-tolerance selection. Three cultivars, Taishan 6426, Taishan 4033, and Jimai 22, were exposed to LT stress at standing, jointing and booting stages, and the yield-related traits, photosynthetic parameters as well as leaf cell structure were compared among different cultivars. The agronomic traits of different cultivars changed in similar trends after LT treatment with the consistent result across years. Plant height received small influence of LT, especially LT at standing stage. In contrast, spike number per plant decreased significantly compared with the control(normal temperature), but the reduced percentage varied across years. Grain number per spike and yield per plant showed significantly declined trends with delaying LT stress, and the maximum decrements were observed in the treatment of LT at booting stage. The grain number per spike was 90.5%–93.3%(2012–2013) and 91.9%–93.6%(2013–2014) lower, and the yield per plant was 87.9%–97.3%(2012–2013) and 91.5%–97.8%(2013–2014) lower in LT treatment at booting than in the control. Booting stage was secondly sensitive to LT in yield, whereas jointing stage was less influenced by LT. Grain yield loss under LT stress mainly resulted from the decrease of grain number per spike. Under LT at jointing or booting stage, the chlorophyll content decreased in the three cultivars, but the decrement varied across cultivars. Moreover, the photosynthetic rate, transpiration rate and stomata conductance decreased significantly and the intercellular CO_2 concentration increased greatly in the three cultivars, and those in Taishan 6426 had the minimum variations. LT also had inhibition in chlorophyll fluorescence parameters of wheat. For example, the F_v/F_m value was significantly lower in the treatment of LT at jointing or booting stage than in the control except for Taishan 6426. When exposed to LT stress, leaf epidermic cells were severely damaged, showing their loose arrangement and irregular structures of guard cell and accessory cell. Our results indicate that photosynthetic rate, transpiration rate, stomata conductance, intercellular CO_2 concentration and F_v/F_m can be used as indicators to identify tolerance of wheat cultivars to spring coldness.
Low temperature(LT) stress in spring occurs frequently in the Huang-Huai Rivers Valley, resulting in yield loss of wheat. The objectives of this study were to understand the physiological responses of wheat cultivars to LT and determine suitable physiological indicator(s) for cold-tolerance selection. Three cultivars, Taishan 6426, Taishan 4033, and Jimai 22, were exposed to LT stress at standing, jointing and booting stages, and the yield-related traits, photosynthetic parameters as well as leaf cell structure were compared among different cultivars. The agronomic traits of different cultivars changed in similar trends after LT treatment with the consistent result across years. Plant height received small influence of LT, especially LT at standing stage. In contrast, spike number per plant decreased significantly compared with the control(normal temperature), but the reduced percentage varied across years. Grain number per spike and yield per plant showed significantly declined trends with delaying LT stress, and the maximum decrements were observed in the treatment of LT at booting stage. The grain number per spike was 90.5%–93.3%(2012–2013) and 91.9%–93.6%(2013–2014) lower, and the yield per plant was 87.9%–97.3%(2012–2013) and 91.5%–97.8%(2013–2014) lower in LT treatment at booting than in the control. Booting stage was secondly sensitive to LT in yield, whereas jointing stage was less influenced by LT. Grain yield loss under LT stress mainly resulted from the decrease of grain number per spike. Under LT at jointing or booting stage, the chlorophyll content decreased in the three cultivars, but the decrement varied across cultivars. Moreover, the photosynthetic rate, transpiration rate and stomata conductance decreased significantly and the intercellular CO_2 concentration increased greatly in the three cultivars, and those in Taishan 6426 had the minimum variations. LT also had inhibition in chlorophyll fluorescence parameters of wheat. For example, the F_v/F_m value was significantly lower in the treatment of LT at jointing or booting stage than in the control except for Taishan 6426. When exposed to LT stress, leaf epidermic cells were severely damaged, showing their loose arrangement and irregular structures of guard cell and accessory cell. Our results indicate that photosynthetic rate, transpiration rate, stomata conductance, intercellular CO_2 concentration and F_v/F_m can be used as indicators to identify tolerance of wheat cultivars to spring coldness.
引文
[1]Fang S B,Yang J J,Zhou G S.Change trend distributive characteristics of agrometeorological disasters in China in recent 30years.J Nat Dis,2011,20:69-73
[2]Wang J F,Zhu Y Y,Liu H P.Main agrometeorological disasters and their impact in recent 28 years in Henan.Meteorol Environ Sci,2007,30:9-11
[3]Cao H M,Shi Z M,Zhou X B,Lei P Z,Dong S G.A review on response of plant to low temperature and its cold resistance.Chin J Agrometeorol,2010,31:310-314
[4]Gana J A,Sutton F,Kenefick D G.c DNA structure and expression patterns of a low-temperature-specific wheat gene tacr7.Plant Mol Biol,1997,34:643-650
[5]Minami A,Nagao M,Ikegami K,Koshiba T,Arakawa K,Fujikawa S,Takezawa D.Cold acclimation in bryophytes:lowtemperature-induced freezing tolerance in Physcomitrella patens is associated with increases in expression levels of stress-related genes but not with increase in level of endogenous abscisic acid.Planta,2005,220:414-423
[6]刘艳阳,李俊周,陈磊,崔党群.低温胁迫对小麦叶片细胞膜脂质过氧化产物及相关酶活性的影响.麦类作物学报,2006,26(4):70-73Liu Y Y,Li J Z,Chen L,Cui D Q.Effect of low temperature stress on peroxidation product of membrane lipids and activity of related enzymes in wheat seeding leaves.J Triticeae Crops,2006,26(4):70-73(in Chinese with English abstract)
[7]姜丽娜,张黛静,宋飞,刘佩,樊婷婷,余海波,李春喜.不同品种小麦叶片对拔节期低温的生理响应及抗寒性评价.生态学报,2014,34:4251-4261Jiang L N,Zhang D J,Song F,Liu P,Fan T T,Yu H B,Li C X.Evaluation of cold resistance of different wheat varieties based on physiological responses of leaves to low temperature at the jointing stage.Acta Ecol Sin,2014,34:4251-4261(in Chinese with English abstract)
[8]张志伟,王法宏,李升东,冯波,司纪升,张宾.不同类型小麦品种孕穗期低温生理反应及其抗寒性分析。麦类作物学报,2012,32:900-906Zhang Z W,Wang F H,Li S D,Feng B,Si J S,Zhang B.Analysis on physiological reaction to low temperature at booting stage and their cold resistance of different types of wheat varieties.JTriticeae Crops,2012,32:900-906(in Chinese with English abstract)
[9]张军,鲁敏,孙树贵,杜万里,刘洋,武军,陈新宏.拔节期低温胁迫对小麦生理生化特性和产量的影响.西北农业学报,2014,23(2):73-79Zhang J,Lu M,Sun S G,Du W L,Liu Y,Wu J,Chen X H.Changes of physiological and biochemical parameters and grain yield at joining stage of wheat under low temperature stress.Acta Agric Boreali-Occident Sin,2014,23(2):73-79(in Chinese with English abstract)
[10]孙金月,赵玉田,梁博文,刘方,尤勇.HRGP在小麦抗寒锻炼过程中的变化及其与抗寒性的关系.植物遗传资源学报,2004,5:6-11Sun J Y,Zhao Y T,Liang B W,Liu F,You Y.Changes of hydroxyproline-rich glyco-protein wheat under cold stress and its relationship to cold resistance.Plant Genet Resour,2004,5:6-11(in Chinese with English abstract)
[11]Equiza M A.Morphological,anatomical and physiological responses related to differential shoot vs.root growth inhibition at low temperature in spring and winter wheat.Ann Bot,2001,87:67-76
[12]刘艳阳,李俊周,陈磊,崔党群.低温胁迫对小麦叶片细胞膜脂质过氧化产物及相关酶活性的影响.麦类作物学报,2006,26(4):70-73Liu Y Y,Li J Z,Chen L,Cui D Q.Effect of low temperature stress on peroxidation product of membrane lipids and activity of related enzymes in wheat seed-ling leaves.J Triticeae Crops,2006,26(4):70-73(in Chinese with English abstract)
[13]Bohn M,Luthje S.Plasma membrane lipid alterations induced by cold acclimation and abscisic acid treatment of winter wheat seedlings differing in frost resistance.J Plant Physiol,2007,164:146-156
[14]刘炜,孙德兰,王红,简令成,张可夫.2?C低温下抗寒冬小麦与冷敏感春小麦幼苗细胞质膜Ca2+-ATPase活性比较.作物学报,2002,28:227-229Liu W,Sun D L,Wang H,Jian L C,Zhao K F.Comparison of Ca2+-ATPase activity in seedling plasmolemma of cold resistant winter wheat vs.cold-sensitive spring wheat under 2?C low temperature.Acta Agron Sin,2002,28:227-229(in Chinese with English abstract)
[15]蒋高明.植物生理生态学.北京:高等教育出版社,2004.pp46-53Jiang G M.Plant Ecophysiology.Beijing:Higher Education Press,2004.pp 46-53(in Chinese)
[16]Genty B,Briantais J M,Baker N R.The relationship between the quantum yield of photosynthetic electron transport and quenching of chlorophyll fluorescence.Biochim Biophys Acta,1989,99:87-92
[17]Schreiber U,Bilger W,Neubauer C.Chlorophyll fluorescence as a non-destructive indicator for rapid assessment of in vivo photosynthesis.Ecol Studies,1994,100:49-70
[18]Krause G H,Weis E.Chlorophyll fluorescence and photosynthesis:the basis.Annu Rev Plant Physiol Plant Mol Biol,1991,42:313-349
[19]Van Kooten O,Snel J F.The use of chlorophyll fluorescence nomenclature in plant stress physiology.Photosynth Res,1990,25:147-150
[20]王晓楠,付连双,李卓夫,杨方,孙艳丽,刘灿,王金伟,陈禹兴.低温驯化及封冻阶段不同冬小麦品种叶绿素荧光参数的比较.麦类作物学报,2009,29:83-88Wang X N,Fu L S,Li Z F,Yang F,Sun Y L,Liu C,Wang J W,Chen Y X.Comparison of chlorophyll fluorescence parameters between two winter wheat cultivars during cold acclimation and frozen period.J Triticeae Crops,2009,29:83-88(in Chinese with English abstract)
[21]高艳,唐建卫,殷贵鸿,韩玉林,黄峰,王丽娜.倒春寒发生时期和次数对冬小麦产量性状的影响.麦类作物学报,2015,35:687-692Gao Y,Tang J W,Yin G H,Han Y L,Huang F,Wang L N.Effect of different periods and frequency of late spring coldness on winter wheat yield related traits.J Triticeae Crops,2015,35:687-692(in Chinese with English abstract)
[22]王树刚,王振林,王平,王海伟,黄玮,武玉国,尹燕枰.基于生理指标与籽粒产量关系的小麦品种抗冻性分析.应用生态学报,2011,22:1477-1484Wang S G,Wang Z L,Wang P,Wang H W,Huang W,Wu Y G,Yin Y P.Freeze resistance analysis of different wheat cultivars based on relationships between physiological indices and grain yield.Chin J Appl Ecol,2011,22:1477-1484(in Chinese with English abstract)
[23]Fracheboud Y,Haldimann P,Leipner J,Stamp P.Chlorophyll fluorescence as a selection tool for cold tolerance of photosynthesis in maize.J Exp Bot,1999,50:1533-1540
[24]Xu C C,Zhang J H.Effect of drought on chlorophyll fluorescence and xanthophylls cycle components in winter wheat leaves with different ages.Acta Phytophysiol Sin,1999,25:29-37
[25]Kratsh H A,Wise R R.The ultrastructure of chilling stress.Plant Cell Environ,2000,23:337-350
[26]陈思思,李春燕,杨景,徐雯,朱新开,郭文善,封超年.拔节期低温冻害对扬麦16光合特性及产量形成的影响.扬州大学学报(农业与生命科学版),2014,35(3):59-64Chen S S,Li C Y,Yang J,Xu W,Zhu X K,Guo W S,Feng C N.Effect of low temperature at jointing stage on photosynthetic characteristics and yield in wheat cultivar Yangmai 16.J Yangzhou Univ(Agric Life Sci Edn),2014,35(3):59-64(in Chinese with English abstract)
[27]简令成.植物抗寒机理研究的新进展.植物学通报,1992,9(3):17-22Jian L C.Advance of the studies on the mechanism of plant cold hardiness.Chin Bull Bot,1992,9(3):17-22(in Chinese with English abstract)
[2]Wang J F,Zhu Y Y,Liu H P.Main agrometeorological disasters and their impact in recent 28 years in Henan.Meteorol Environ Sci,2007,30:9-11
[3]Cao H M,Shi Z M,Zhou X B,Lei P Z,Dong S G.A review on response of plant to low temperature and its cold resistance.Chin J Agrometeorol,2010,31:310-314
[4]Gana J A,Sutton F,Kenefick D G.c DNA structure and expression patterns of a low-temperature-specific wheat gene tacr7.Plant Mol Biol,1997,34:643-650
[5]Minami A,Nagao M,Ikegami K,Koshiba T,Arakawa K,Fujikawa S,Takezawa D.Cold acclimation in bryophytes:lowtemperature-induced freezing tolerance in Physcomitrella patens is associated with increases in expression levels of stress-related genes but not with increase in level of endogenous abscisic acid.Planta,2005,220:414-423
[6]刘艳阳,李俊周,陈磊,崔党群.低温胁迫对小麦叶片细胞膜脂质过氧化产物及相关酶活性的影响.麦类作物学报,2006,26(4):70-73Liu Y Y,Li J Z,Chen L,Cui D Q.Effect of low temperature stress on peroxidation product of membrane lipids and activity of related enzymes in wheat seeding leaves.J Triticeae Crops,2006,26(4):70-73(in Chinese with English abstract)
[7]姜丽娜,张黛静,宋飞,刘佩,樊婷婷,余海波,李春喜.不同品种小麦叶片对拔节期低温的生理响应及抗寒性评价.生态学报,2014,34:4251-4261Jiang L N,Zhang D J,Song F,Liu P,Fan T T,Yu H B,Li C X.Evaluation of cold resistance of different wheat varieties based on physiological responses of leaves to low temperature at the jointing stage.Acta Ecol Sin,2014,34:4251-4261(in Chinese with English abstract)
[8]张志伟,王法宏,李升东,冯波,司纪升,张宾.不同类型小麦品种孕穗期低温生理反应及其抗寒性分析。麦类作物学报,2012,32:900-906Zhang Z W,Wang F H,Li S D,Feng B,Si J S,Zhang B.Analysis on physiological reaction to low temperature at booting stage and their cold resistance of different types of wheat varieties.JTriticeae Crops,2012,32:900-906(in Chinese with English abstract)
[9]张军,鲁敏,孙树贵,杜万里,刘洋,武军,陈新宏.拔节期低温胁迫对小麦生理生化特性和产量的影响.西北农业学报,2014,23(2):73-79Zhang J,Lu M,Sun S G,Du W L,Liu Y,Wu J,Chen X H.Changes of physiological and biochemical parameters and grain yield at joining stage of wheat under low temperature stress.Acta Agric Boreali-Occident Sin,2014,23(2):73-79(in Chinese with English abstract)
[10]孙金月,赵玉田,梁博文,刘方,尤勇.HRGP在小麦抗寒锻炼过程中的变化及其与抗寒性的关系.植物遗传资源学报,2004,5:6-11Sun J Y,Zhao Y T,Liang B W,Liu F,You Y.Changes of hydroxyproline-rich glyco-protein wheat under cold stress and its relationship to cold resistance.Plant Genet Resour,2004,5:6-11(in Chinese with English abstract)
[11]Equiza M A.Morphological,anatomical and physiological responses related to differential shoot vs.root growth inhibition at low temperature in spring and winter wheat.Ann Bot,2001,87:67-76
[12]刘艳阳,李俊周,陈磊,崔党群.低温胁迫对小麦叶片细胞膜脂质过氧化产物及相关酶活性的影响.麦类作物学报,2006,26(4):70-73Liu Y Y,Li J Z,Chen L,Cui D Q.Effect of low temperature stress on peroxidation product of membrane lipids and activity of related enzymes in wheat seed-ling leaves.J Triticeae Crops,2006,26(4):70-73(in Chinese with English abstract)
[13]Bohn M,Luthje S.Plasma membrane lipid alterations induced by cold acclimation and abscisic acid treatment of winter wheat seedlings differing in frost resistance.J Plant Physiol,2007,164:146-156
[14]刘炜,孙德兰,王红,简令成,张可夫.2?C低温下抗寒冬小麦与冷敏感春小麦幼苗细胞质膜Ca2+-ATPase活性比较.作物学报,2002,28:227-229Liu W,Sun D L,Wang H,Jian L C,Zhao K F.Comparison of Ca2+-ATPase activity in seedling plasmolemma of cold resistant winter wheat vs.cold-sensitive spring wheat under 2?C low temperature.Acta Agron Sin,2002,28:227-229(in Chinese with English abstract)
[15]蒋高明.植物生理生态学.北京:高等教育出版社,2004.pp46-53Jiang G M.Plant Ecophysiology.Beijing:Higher Education Press,2004.pp 46-53(in Chinese)
[16]Genty B,Briantais J M,Baker N R.The relationship between the quantum yield of photosynthetic electron transport and quenching of chlorophyll fluorescence.Biochim Biophys Acta,1989,99:87-92
[17]Schreiber U,Bilger W,Neubauer C.Chlorophyll fluorescence as a non-destructive indicator for rapid assessment of in vivo photosynthesis.Ecol Studies,1994,100:49-70
[18]Krause G H,Weis E.Chlorophyll fluorescence and photosynthesis:the basis.Annu Rev Plant Physiol Plant Mol Biol,1991,42:313-349
[19]Van Kooten O,Snel J F.The use of chlorophyll fluorescence nomenclature in plant stress physiology.Photosynth Res,1990,25:147-150
[20]王晓楠,付连双,李卓夫,杨方,孙艳丽,刘灿,王金伟,陈禹兴.低温驯化及封冻阶段不同冬小麦品种叶绿素荧光参数的比较.麦类作物学报,2009,29:83-88Wang X N,Fu L S,Li Z F,Yang F,Sun Y L,Liu C,Wang J W,Chen Y X.Comparison of chlorophyll fluorescence parameters between two winter wheat cultivars during cold acclimation and frozen period.J Triticeae Crops,2009,29:83-88(in Chinese with English abstract)
[21]高艳,唐建卫,殷贵鸿,韩玉林,黄峰,王丽娜.倒春寒发生时期和次数对冬小麦产量性状的影响.麦类作物学报,2015,35:687-692Gao Y,Tang J W,Yin G H,Han Y L,Huang F,Wang L N.Effect of different periods and frequency of late spring coldness on winter wheat yield related traits.J Triticeae Crops,2015,35:687-692(in Chinese with English abstract)
[22]王树刚,王振林,王平,王海伟,黄玮,武玉国,尹燕枰.基于生理指标与籽粒产量关系的小麦品种抗冻性分析.应用生态学报,2011,22:1477-1484Wang S G,Wang Z L,Wang P,Wang H W,Huang W,Wu Y G,Yin Y P.Freeze resistance analysis of different wheat cultivars based on relationships between physiological indices and grain yield.Chin J Appl Ecol,2011,22:1477-1484(in Chinese with English abstract)
[23]Fracheboud Y,Haldimann P,Leipner J,Stamp P.Chlorophyll fluorescence as a selection tool for cold tolerance of photosynthesis in maize.J Exp Bot,1999,50:1533-1540
[24]Xu C C,Zhang J H.Effect of drought on chlorophyll fluorescence and xanthophylls cycle components in winter wheat leaves with different ages.Acta Phytophysiol Sin,1999,25:29-37
[25]Kratsh H A,Wise R R.The ultrastructure of chilling stress.Plant Cell Environ,2000,23:337-350
[26]陈思思,李春燕,杨景,徐雯,朱新开,郭文善,封超年.拔节期低温冻害对扬麦16光合特性及产量形成的影响.扬州大学学报(农业与生命科学版),2014,35(3):59-64Chen S S,Li C Y,Yang J,Xu W,Zhu X K,Guo W S,Feng C N.Effect of low temperature at jointing stage on photosynthetic characteristics and yield in wheat cultivar Yangmai 16.J Yangzhou Univ(Agric Life Sci Edn),2014,35(3):59-64(in Chinese with English abstract)
[27]简令成.植物抗寒机理研究的新进展.植物学通报,1992,9(3):17-22Jian L C.Advance of the studies on the mechanism of plant cold hardiness.Chin Bull Bot,1992,9(3):17-22(in Chinese with English abstract)