白三叶不同叶龄叶片对不同季节温度适应的生理调控机理
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摘要
对不同季节白三叶(Trifolium repens)不同叶龄叶片(幼叶、中叶和老叶)抗氧化酶(SOD、POD和CAT)活力、渗透调节物(可溶性糖、可溶性蛋白和游离脯氨酸)含量、MDA含量和叶绿素含量进行分析,揭示白三叶不同叶龄叶片对不同季节温度适应的生理调控机理和白三叶叶片短寿在维护匍匐茎生长中的生态作用。结果表明:(1)冬季低温生出的幼叶MDA含量较低,而叶绿素a、POD和SOD活力、可溶性糖和脯氨酸含量均最高;夏季高温生出的幼叶MDA和叶绿素b含量较高,而POD和SOD活力、可溶性糖和脯氨酸含量较低。白三叶幼叶具有较强的生理调控可塑性以适应冬夏季温度。(2)不同季节幼叶成长过程中,其生理调控模式相近,随叶片长大成熟,叶绿素含量和SOD、POD、CAT活力及脯氨酸和可溶性糖含量均增高,而MDA含量降低。(3)不同季节老化叶片生理特征相同,叶片中可溶性糖和脯氨酸含量趋于增高,CAT活力及MDA含量均下降。研究表明,抗氧化酶和渗透调节物通过维护氧自由基代谢和水分代谢平衡而在幼叶适应不同季节温度、叶片快速生长中起重要保护作用。不同季节叶片抗氧化能力下降和持续积累的脯氨酸、可溶性糖抑制光合作用可能是叶片衰老和短寿的诱因。不同季节匍匐茎上叶片的短寿、快速衰老可使匍匐茎将能量物质分配给其顶部,促进匍匐茎顶部幼叶萌生和匍匐茎的延伸生长及种群的扩散,其在维护白三叶植株持续生存中具有重要的调控作用。
In this study,the activities of superoxide dismutase( SOD),peroxide( POD) and catalse( CAT),and the contents of soluble sugar,soluble protein,proline,malondialdehyde and chlorophyll in different aged leaves( young,middle-aged,old) of white clover( Trifolium repens) were investigated in different seasons to understand how different aged leaves of white clover adapt to different seasonal temperatures in physiology and what ecological role of leaf senescence plays in maintaining stolon growth. The result showed that:( 1) young leaves grown in winter had lower MDA content,and higher values in chlorophyll a content,SOD and POD activities,and soluble sugar and proline contents. However,young leaves grown in summer had higher MDA and chlorophyll b contents,but had lower SOD and POD activities and soluble sugar and proline contents. It is indicated that young leaves had higher physiological regulation plasticity in adaptation to different seasonal temperatures.( 2) There existed a similar model of physiological regulation in the developing leaves grown in different seasons. When young leaves became mature,the contents of chlorophyll and soluble sugar and proline,and the activities of SOD and POD increased,while MDA content decreased.( 3) Aging leaves grown in different seasons had same physiological characteristics. With the leaves becoming senescent,the soluble sugar and proline contents still increased,but the activities of SOD and CAT and content of MDA decreased. The antioxidant enzyme system and osmoregulation played an important role in physiological protection of young leaves to adapt different seasonal temperatures and in keeping their rapid growth by maintaining metabolic balance of oxygen free radicals and water metabolic balance.During leaf development,the antioxidant capacity decreased,and the proline and soluble sugar kept on accumulation,which was the reason inducing leaf senescence and short life. However,short life of leaves grown in different seasons would make more energy and matter being allocated to stolon top and promote leaf initiation,stolon elongation growth,and population expansion,which played an important physiological regulating role in maintaining growth of white clover in different seasons.
In this study,the activities of superoxide dismutase( SOD),peroxide( POD) and catalse( CAT),and the contents of soluble sugar,soluble protein,proline,malondialdehyde and chlorophyll in different aged leaves( young,middle-aged,old) of white clover( Trifolium repens) were investigated in different seasons to understand how different aged leaves of white clover adapt to different seasonal temperatures in physiology and what ecological role of leaf senescence plays in maintaining stolon growth. The result showed that:( 1) young leaves grown in winter had lower MDA content,and higher values in chlorophyll a content,SOD and POD activities,and soluble sugar and proline contents. However,young leaves grown in summer had higher MDA and chlorophyll b contents,but had lower SOD and POD activities and soluble sugar and proline contents. It is indicated that young leaves had higher physiological regulation plasticity in adaptation to different seasonal temperatures.( 2) There existed a similar model of physiological regulation in the developing leaves grown in different seasons. When young leaves became mature,the contents of chlorophyll and soluble sugar and proline,and the activities of SOD and POD increased,while MDA content decreased.( 3) Aging leaves grown in different seasons had same physiological characteristics. With the leaves becoming senescent,the soluble sugar and proline contents still increased,but the activities of SOD and CAT and content of MDA decreased. The antioxidant enzyme system and osmoregulation played an important role in physiological protection of young leaves to adapt different seasonal temperatures and in keeping their rapid growth by maintaining metabolic balance of oxygen free radicals and water metabolic balance.During leaf development,the antioxidant capacity decreased,and the proline and soluble sugar kept on accumulation,which was the reason inducing leaf senescence and short life. However,short life of leaves grown in different seasons would make more energy and matter being allocated to stolon top and promote leaf initiation,stolon elongation growth,and population expansion,which played an important physiological regulating role in maintaining growth of white clover in different seasons.
引文
安莹,陈雅君,赵伟,等.2009.低温对白三叶膜质过氧化及保护酶活性的影响.草原与草坪,134(3):8-10.
范玉贞.2010.自然低温胁迫对白三叶草的膜损伤与SOD活性的影响.湖北农业科学,49(2):404-409.
高婷婷,毛培春,田小霞.2014.10个白三叶品种苗期耐寒性评价.草业科学,31(9):1724-1731.
和红云,薛琳,田丽萍,等.2008.低温胁迫对甜瓜幼苗叶绿素含量及荧光参数的影响.北方园艺,(4):13-16.
李州,彭燕,苏星源.2013.不同叶型白三叶抗氧化保护及渗透调节生理对干旱胁迫的响应.草业学报,22(2):257-263.
梁超,王晓雷.2013.浅析白三叶在辽西地区园林绿化中的应用.现代园林,(3):111-113
林植芳,刘楠.2012.活性氧调控植物生长发育的研究进展.植物学报,47(1):74-86.
张盼盼,冯佰利,王鹏科,等.2010.干旱条件下糜子叶片衰老与保护酶活性变化.干旱地区农业研究,28(2):99-103.
张志良,瞿伟菁.2003.植物生理学实验指导(第三版).北京:高等教育出版社.
赵梅,周瑞莲,刘建芳,等.2011.冬季融冻过程中白三叶叶片抗氧化酶活力和渗透调节物含量变化与抗冻性相关关系.生态学报,31(2):306-315.
周瑞莲,赵梅,王进,等.2012a.冬季高温对白三叶越冬和适应春季“倒春寒”的影响.生态学报,32(14):4462-4471.
周瑞莲,赵梅,张萍,等.2012b.白三叶和红三叶对人工融冻-冻融胁迫生理响应的差异.生态学杂志,31(6):1334-1340.
Bajji M,Lutts S,Kinet JM.2001.Water deficit effects on solute contribution to osmotic adjustment as a function of leaf ageing in three durum wheat(Triticum durum Desf.)cultivars performing differently in arid conditions.Plant Science,160:669-681.
Drazkiewicz M,Skórzyńska-Polit E,Krupa Z.2004.Copper-induced oxidative stress and antioxidant defence in Arabidopsis thaliana.Biometals,17:379-387.
Ewers FW,Lawson MC,Bowen TJ,et al.2003.Freeze/thaw stress in Ceanothus of southern California chaparral.Oecologia,136:213-219.
Guinchard MP,Robin CH,Grieu PH.1997.Cold acclimation in white clover subjected to chilling and frost:Changes in water and carbohydrates status.European Journal of Agronomy,6:225-233.
Kameli A,Losel DM.1995.Contribution of carbohydrates and other solutes to osmotic adjustment in wheat leaves under water stress.Journal of Plant Physiology,145:363-366.
Kirsti R,OLavi J,Arild E,et al.1993.Development of cold tolerance in white clover(Trifolium repens L.)in relation to carbohydrate and free amino acid content.Acta Agriculturae Scandinavica,43:151-155.
Lee BR,Li LS,Jung WJ,et al.2009.Water deficit-induced oxidative stress and the activation of antioxidant enzymes in white clover leaves.Biologia Plantarum,53:505-510.
Ono K,Nishi Y,Watanabe A,et al.2001.Possible mechanisms of adaptive leaf senescence.Plant Biology,3:234-243.
Rolland F,Moore B,Sheen J.2002.Sugar sensing and signaling in plants.The Plant Cell,14(Suppl):185-205.
Shimazake K,Sakaki T,Kondo N,et al.1980.Active oxygen participation in chlorophyll destruction and lipid peroxidation in SO2fumigated leaves of spinach.Plant Cell Physiology,21:1193-1204.
Sundar D,Perianayaguy B,Reddy AR.2004.Localization of antioxidant enzymes in the cellular compartments of sorghum leaves.Plant Growth Regulation,44:157-163.
Yoshida S.2003.Molecular regulation of leaf senescence.Current Opinion in Plant Biology,6:79-84.
范玉贞.2010.自然低温胁迫对白三叶草的膜损伤与SOD活性的影响.湖北农业科学,49(2):404-409.
高婷婷,毛培春,田小霞.2014.10个白三叶品种苗期耐寒性评价.草业科学,31(9):1724-1731.
和红云,薛琳,田丽萍,等.2008.低温胁迫对甜瓜幼苗叶绿素含量及荧光参数的影响.北方园艺,(4):13-16.
李州,彭燕,苏星源.2013.不同叶型白三叶抗氧化保护及渗透调节生理对干旱胁迫的响应.草业学报,22(2):257-263.
梁超,王晓雷.2013.浅析白三叶在辽西地区园林绿化中的应用.现代园林,(3):111-113
林植芳,刘楠.2012.活性氧调控植物生长发育的研究进展.植物学报,47(1):74-86.
张盼盼,冯佰利,王鹏科,等.2010.干旱条件下糜子叶片衰老与保护酶活性变化.干旱地区农业研究,28(2):99-103.
张志良,瞿伟菁.2003.植物生理学实验指导(第三版).北京:高等教育出版社.
赵梅,周瑞莲,刘建芳,等.2011.冬季融冻过程中白三叶叶片抗氧化酶活力和渗透调节物含量变化与抗冻性相关关系.生态学报,31(2):306-315.
周瑞莲,赵梅,王进,等.2012a.冬季高温对白三叶越冬和适应春季“倒春寒”的影响.生态学报,32(14):4462-4471.
周瑞莲,赵梅,张萍,等.2012b.白三叶和红三叶对人工融冻-冻融胁迫生理响应的差异.生态学杂志,31(6):1334-1340.
Bajji M,Lutts S,Kinet JM.2001.Water deficit effects on solute contribution to osmotic adjustment as a function of leaf ageing in three durum wheat(Triticum durum Desf.)cultivars performing differently in arid conditions.Plant Science,160:669-681.
Drazkiewicz M,Skórzyńska-Polit E,Krupa Z.2004.Copper-induced oxidative stress and antioxidant defence in Arabidopsis thaliana.Biometals,17:379-387.
Ewers FW,Lawson MC,Bowen TJ,et al.2003.Freeze/thaw stress in Ceanothus of southern California chaparral.Oecologia,136:213-219.
Guinchard MP,Robin CH,Grieu PH.1997.Cold acclimation in white clover subjected to chilling and frost:Changes in water and carbohydrates status.European Journal of Agronomy,6:225-233.
Kameli A,Losel DM.1995.Contribution of carbohydrates and other solutes to osmotic adjustment in wheat leaves under water stress.Journal of Plant Physiology,145:363-366.
Kirsti R,OLavi J,Arild E,et al.1993.Development of cold tolerance in white clover(Trifolium repens L.)in relation to carbohydrate and free amino acid content.Acta Agriculturae Scandinavica,43:151-155.
Lee BR,Li LS,Jung WJ,et al.2009.Water deficit-induced oxidative stress and the activation of antioxidant enzymes in white clover leaves.Biologia Plantarum,53:505-510.
Ono K,Nishi Y,Watanabe A,et al.2001.Possible mechanisms of adaptive leaf senescence.Plant Biology,3:234-243.
Rolland F,Moore B,Sheen J.2002.Sugar sensing and signaling in plants.The Plant Cell,14(Suppl):185-205.
Shimazake K,Sakaki T,Kondo N,et al.1980.Active oxygen participation in chlorophyll destruction and lipid peroxidation in SO2fumigated leaves of spinach.Plant Cell Physiology,21:1193-1204.
Sundar D,Perianayaguy B,Reddy AR.2004.Localization of antioxidant enzymes in the cellular compartments of sorghum leaves.Plant Growth Regulation,44:157-163.
Yoshida S.2003.Molecular regulation of leaf senescence.Current Opinion in Plant Biology,6:79-84.