脱落酸和乙烯对水稻与小麦籽粒灌浆的调控作用及其机理
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摘要
脱落酸(ABA)和乙烯是两类重要的植物激素,对植物生长发育特别是对逆境响应起重要调控作用。但有关ABA和乙烯对稻麦籽粒灌浆的作用及其机理,目前尚不清楚。本研究以代表性水稻和小麦品种为材料,研究了ABA和乙烯对水稻与小麦籽粒灌浆的调控作用及其机理。主要结果如下:
1. ABA和乙烯对稻麦胚乳发育和充实的调控水稻或小麦穗上强势粒胚乳细胞增殖速率、胚乳细胞数目、籽粒灌浆速率及粒重都高于弱势粒。灌浆初期强势粒乙烯释放速率和1-氨基环丙烷-1-羧酸(ACC)浓度则低于弱势粒,强势粒ABA浓度及ABA/ACC则显著高于弱势粒。胚乳细胞增殖速率和籽粒灌浆速率与乙烯释放速率呈极显著的负相关,与ABA含量及ABA/ACC呈极显著正相关。于灌浆初期(花后0～6天)喷施硝酸钴(乙烯合成抑制剂)或ABA,显著降低了弱势籽粒乙烯释放速率和ACC浓度,增加了胚乳细胞分裂速率、最大细胞数目、籽粒灌浆速率和粒重;喷施乙烯利(乙烯释放促进剂)或氟草酮(ABA合成抑制剂),结果则相反。表明ABA和乙烯及其相互作用对稻麦胚乳细胞增殖和籽粒灌浆起重要调控作用。较高的ABA与乙烯(ACC)的比值有利于稻麦胚乳细胞发育和籽粒灌浆。
2. ABA和乙烯对土壤水分的响应及其与稻麦籽粒灌浆的关系在稻麦灌浆期进行土壤轻度落干或轻度干旱(MD)和土壤重度落干或重度干旱(SD)处理,以土壤水分充足(WW)为对照,研究了ABA和乙烯对土壤水分的响应及其与籽粒灌浆的关系。结果表明,与WW处理相比,MD处理显著提高了籽粒灌浆速率和粒重,SD处理明显降低了籽粒灌浆速率和粒重。在各土壤水分状况下,籽粒乙烯的释放速率和ACC含量在灌浆早期很高,随灌浆天数的增加而迅速降低。籽粒ABA含量在灌浆初期较低,随籽粒生育进程而不断增加,达到最大值后下降。籽粒中ABA浓度随土壤水分的降低而显著增加。籽粒中乙烯释放速率和ACC浓度在MD处理下降低,在SD处理下增加。MD增加了籽粒中ABA与ACC的比值,SD则降低了ABA/ACC值。籽粒灌浆速率与籽粒ABA浓度及ABA/ACC值呈极显著正相关,与乙烯释放速率呈极显著负相关。在花后9～13天喷施乙烯合成抑制物质氨基-乙氧基乙烯基甘氨酸(AVG)或硝酸钴或ABA明显地降低了籽粒ACC浓度和乙烯的释放速率,显著提高了籽粒灌浆速率和粒重以及籽粒中蔗糖合成酶(SuSase)、腺苷二磷酸葡萄糖焦磷酸化酶(AGPase)和可溶性淀粉合成酶(SSSase)活性;喷施乙烯释放促进物质乙烯利或ABA合成抑制物质氟草酮,结果则相反。表明结实期土壤轻度落干或轻度干旱处理可以增加籽粒中ABA与乙烯(ACC)的比值,促进籽粒灌浆。籽粒中ABA和乙烯通过调控籽粒中蔗糖-淀粉代谢途径关键酶活性,实现对籽粒灌浆的调控。
3.不同土壤水分条件下稻麦籽粒ABA和乙烯与膜脂过氧化和核酸含量/浓度的关系籽粒中过氧化氢酶(CAT)和超氧化物岐化酶(SOD)活性在MD处理下显著提高,在SD处理下CAT和SOD的活性在灌浆前期显著提高,在灌浆后期则显著降低。灌浆前期弱势粒中CAT、过氧化物酶(POD)和SOD活性低于强势粒,且受土壤水分的影响要大于强势粒。MD处理籽粒中超氧阴离子生成速率和丙二醛(MDA)含量略有提高,但差异不显著;SD处理显著提高了籽粒中超氧阴离子生成速率和MDA含量。MD处理增加了籽粒核糖核酸(RNA)含量;SD处理则相反。土壤水分对籽粒脱氧核糖核酸(DNA)含量无显著影响。籽粒ABA含量及籽粒灌浆速率与籽粒中SOD和CAT活性及RNA含量呈极显著或显著正相关,与籽粒中MDA含量呈显著负相关。籽粒乙烯释放速率及ACC含量与籽粒中SOD和CAT活性及RNA每粒含量呈显著负相关,与籽粒中超氧阴离子生成速率呈显著正相关。在花后9～13天喷施硝酸钴或ABA,显著地提高了弱势籽粒中CAT和SOD活性,降低了超氧阴离子生成速率和花后18天MDA含量,增加了RNA和DNA含量;喷施乙烯利或氟草酮,结果相反。表明在MD处理下较高的ABA与乙烯(ACC)比值提高了膜脂过氧化保护酶(CAT、SOD)活性,增加了清除自由基(超氧阴离子)的能力,最终促进弱势粒灌浆。
4. ABA和乙烯对稻米品质的调控作用MD处理显著提高了稻米的崩解值,显著降低了垩白粒率、垩白度和消减值;SD处理导致品质变劣。乙烯释放速率与整精米率及崩解值呈负相关,与垩白米率、垩白度及消减值呈正相关;ABA/ACC与整精米率和崩解值呈正相关,与垩白度呈负相关。在花后9～13天喷施乙烯合成抑制物质硝酸钴或ABA,显著提高了整精米率、直链淀粉含量和崩解值,显著降低了稻米的垩白米率、垩白度和蛋白质含量及消减值;喷施乙烯释放促进物质乙烯利或ABA合成抑制物质氟草酮,结果则相反。表明籽粒中ABA和乙烯对水稻品质的形成起调控作用,籽粒中较高的ABA与乙烯比值有利于改善稻米品质。
Both abscisic acid (ABA) and ethylene are important hormones and play important roles in growth and development in responses to stress. It is little known, however, whether and how ABA and ethylene are involved in regulation to grain-filling in rice (Oryza sativa) and wheat (Triticum aestivum). This study investigated the roles of ABA and ethylene in grain filling in rice and wheat and their physiological mechanisms by using typical rice and wheat cultivars. The main results are as follows:
1. Regulation of ABA and ethylene to endosperm development and grain filling. The division rate and the number of endosperm cells, grain filling rate, and grain weight of superior grains in rice or wheat were greater than those of inferior grains. At the early grain filling stage, the evolution rate of ethylene and 1-aminocylopropane-1-carboxylic acid (ACC) concentrations were greater in inferior grains than in superior grains. While ABA concentrations and the ratio of ABA to ACC (ABA/ACC) were greater in the superior than in the inferior grains. The cell division and grain-filling rates were significantly and positively correlated with both ABA contents and the ratio of ABA to ACC, whereas significantly and negatively correlated with ethylene evolution rate. Application of cobalt ion (inhibitor of ethylene synthesis) or ABA at early grain filling stage (at 9-13d post-anthesis) significantly increased endosperm cell division rate and cell number, grain-filling rate and grain weight of inferior spikelets. Application of ethephon (ethylene-releasing agent) or fluridone (inhibitor of carotenoid synthesis) had the opposite effects. The results suggest that the antagonistic interactions between ABA and ethylene mediate the endosperm cell division and grain filling in rice and wheat. A higher ratio of ABA to ethylene (ACC) in rice and wheat grains is required to maintain a faster the endosperm cell division rate and grain-filling rate.
2. Responses of ABA and ethylene in rice and wheat grains to soil moisture and their relations with grain filling. Both moderate soil-drying (MD) and severe soil-drying (SD) were conducted during the grain filling period, and the treatment of well watered (WW) was taken as control. The responses of ABA and ethylene to soil drying and their relations with grain filling were studied. The results showed that, compared with WW, MD significantly increased, whereas SD significantly reduced, grain-filling rate and grain weight. ABA content in the grains was very low at the early grain filling stage, and reached a maximum when the grain-filling rate was the highest. In contrast to ABA, levels of ethylene and ACC in the grains were very high at early grain filling stage and sharply decreased during the linear period of grain growth. Both MD and SD remarkably enhanced ABA accumulation in the grains, which was enhanced more in SD grains than in MD ones. MD reduced, whereas SD remarkably increased, ethylene evolution rate and ACC content. MD increased, whereas SD reduced, the ratio of ABA to ACC. The grain-filling rates were significantly and positively correlated with both ABA contents and the ratio of ABA to ACC, whereas significantly and negatively correlated with ethylene evolution rate. Application of amino-ethoxyvinylglycine (AVG) or cobaltion (inhibitor of ethylene synthesis) or ABA at 9-13 d post-anthesis significantly reduced ACC concentrations and ethylene evolution rate of grains, and significantly enhanced the activities of sucrose synthase (SuSase), ADP glucose pyrophosphorylase (AGPase) and soluble starch synthase (SSSase), and increased grain-filling rate and grain weight. The results were reversed when ethephon (ethylene-releasing agent) or fluridone (inhibitor of ABA synthesis) was applied. The results suggested that a moderate soil-drying during the grain-filling period of rice and wheat could increase the ratio of ABA to ethylene (ACC), and therefore accelerate grain filling and increase grain weight. The mediation of ABA and ethylene to grain filling is probably through regulating key enzymes involved in sucrose-to-starch pathway, and a high ratio of ABA to ACC enhances grain filling.
3. ABA and ethylene in relation with membrane lipid peroxidation and nucleic acid content/concentration in grains at different soil moisture conditions. The MD increased activities of superoxide dismutase (SOD) and catalase (CAT). SD significantly increased those at the early treatment stage, but decreased at the late treatment stage. Among all the treatments, inferior grains had much lower activities of SOD, peroxidase (POD) and CAT than superior grains at the early grain filling stage, and the effect of soil moisture on inferior grains was much grater than on superior grains. There were no significant differences in superoxide anion (O2-) production rate and malondialdehyde (MDA) content of grains between MD and WW plants, but they were much higher in SD plants. MD significantly increased, whereas SD significantly reduced, ribonucleic acid (RNA) contents in grains. There were no significant differences in deoxyribonucleic acid (DNA) contents in grains among all the treatments. The grain-filling rates and ABA contents in grains were significantly and positively correlated with activities of SOD and CAT and RNA contents in grains, whereas significantly and negatively correlated with MDA content of grains. The ethylene evolution rate and ACC content in grains were significantly and negatively correlated with activities of SOD and CAT and RNA contents, and significantly and positively correlated with O2- production rate in grains. Application of cobaltion (inhibitor of ethylene synthesis) or ABA, at 9-13 d post-anthesis significantly enhanced the activities of SOD and CAT, increased RNA and DNA contents of grains, whereas significantly reduced O2- production rate in grains and at MDA content of grains, and the results were reversed when ethephon (ethylene-releasing agent) or fluridone (inhibitor of ABA synthesis) was applied. The results suggested that a moderate soil drying during the grain-filling period of rice and wheat could enhance the ratio of ABA to ethylene (ACC) and the activities of SOD and CAT of grains, and reduce the ethylene evolution rate and membrane lipid peroxidation in grains, and therefore accelerate the filling of inferior grains.
4. Roles of ABA and ethylene in grain quality of rice. The MD significantly increased breakdown values, and reduced chalky kernels, chalkiness and setback values. The results were reversed for the SD treatment. Ethylene evolution rates were significantly and negatively correlated with head milled rice and breakdown values, and significantly and positively correlated with chalky kernels, chalkiness and setback values. The ratio of ABA to ACC was significantly and positively correlated with head milled rice and breakdown values, and significantly and negatively correlated with chalkiness. Application of cobaltion (inhibitor of ethylene synthesis) or ABA, at 9-13 d post-anthesis significantly increased head milled rice, amylose contents and breakdown values, whereas significantly reduced chalky kernels, chalkiness, protein contents, and setback values, the results were reversed when ethephon (ethylene-releasing agent) or fluridone (inhibitor of ABA synthesis) was applied. The results indicate that Both ABA and ethylene play roles in the formation of grain quality, and that a higher ratio of ABA to ethylene (ACC) would benefit grain quality of rice.
1. ABA和乙烯对稻麦胚乳发育和充实的调控水稻或小麦穗上强势粒胚乳细胞增殖速率、胚乳细胞数目、籽粒灌浆速率及粒重都高于弱势粒。灌浆初期强势粒乙烯释放速率和1-氨基环丙烷-1-羧酸(ACC)浓度则低于弱势粒,强势粒ABA浓度及ABA/ACC则显著高于弱势粒。胚乳细胞增殖速率和籽粒灌浆速率与乙烯释放速率呈极显著的负相关,与ABA含量及ABA/ACC呈极显著正相关。于灌浆初期(花后0～6天)喷施硝酸钴(乙烯合成抑制剂)或ABA,显著降低了弱势籽粒乙烯释放速率和ACC浓度,增加了胚乳细胞分裂速率、最大细胞数目、籽粒灌浆速率和粒重;喷施乙烯利(乙烯释放促进剂)或氟草酮(ABA合成抑制剂),结果则相反。表明ABA和乙烯及其相互作用对稻麦胚乳细胞增殖和籽粒灌浆起重要调控作用。较高的ABA与乙烯(ACC)的比值有利于稻麦胚乳细胞发育和籽粒灌浆。
2. ABA和乙烯对土壤水分的响应及其与稻麦籽粒灌浆的关系在稻麦灌浆期进行土壤轻度落干或轻度干旱(MD)和土壤重度落干或重度干旱(SD)处理,以土壤水分充足(WW)为对照,研究了ABA和乙烯对土壤水分的响应及其与籽粒灌浆的关系。结果表明,与WW处理相比,MD处理显著提高了籽粒灌浆速率和粒重,SD处理明显降低了籽粒灌浆速率和粒重。在各土壤水分状况下,籽粒乙烯的释放速率和ACC含量在灌浆早期很高,随灌浆天数的增加而迅速降低。籽粒ABA含量在灌浆初期较低,随籽粒生育进程而不断增加,达到最大值后下降。籽粒中ABA浓度随土壤水分的降低而显著增加。籽粒中乙烯释放速率和ACC浓度在MD处理下降低,在SD处理下增加。MD增加了籽粒中ABA与ACC的比值,SD则降低了ABA/ACC值。籽粒灌浆速率与籽粒ABA浓度及ABA/ACC值呈极显著正相关,与乙烯释放速率呈极显著负相关。在花后9～13天喷施乙烯合成抑制物质氨基-乙氧基乙烯基甘氨酸(AVG)或硝酸钴或ABA明显地降低了籽粒ACC浓度和乙烯的释放速率,显著提高了籽粒灌浆速率和粒重以及籽粒中蔗糖合成酶(SuSase)、腺苷二磷酸葡萄糖焦磷酸化酶(AGPase)和可溶性淀粉合成酶(SSSase)活性;喷施乙烯释放促进物质乙烯利或ABA合成抑制物质氟草酮,结果则相反。表明结实期土壤轻度落干或轻度干旱处理可以增加籽粒中ABA与乙烯(ACC)的比值,促进籽粒灌浆。籽粒中ABA和乙烯通过调控籽粒中蔗糖-淀粉代谢途径关键酶活性,实现对籽粒灌浆的调控。
3.不同土壤水分条件下稻麦籽粒ABA和乙烯与膜脂过氧化和核酸含量/浓度的关系籽粒中过氧化氢酶(CAT)和超氧化物岐化酶(SOD)活性在MD处理下显著提高,在SD处理下CAT和SOD的活性在灌浆前期显著提高,在灌浆后期则显著降低。灌浆前期弱势粒中CAT、过氧化物酶(POD)和SOD活性低于强势粒,且受土壤水分的影响要大于强势粒。MD处理籽粒中超氧阴离子生成速率和丙二醛(MDA)含量略有提高,但差异不显著;SD处理显著提高了籽粒中超氧阴离子生成速率和MDA含量。MD处理增加了籽粒核糖核酸(RNA)含量;SD处理则相反。土壤水分对籽粒脱氧核糖核酸(DNA)含量无显著影响。籽粒ABA含量及籽粒灌浆速率与籽粒中SOD和CAT活性及RNA含量呈极显著或显著正相关,与籽粒中MDA含量呈显著负相关。籽粒乙烯释放速率及ACC含量与籽粒中SOD和CAT活性及RNA每粒含量呈显著负相关,与籽粒中超氧阴离子生成速率呈显著正相关。在花后9～13天喷施硝酸钴或ABA,显著地提高了弱势籽粒中CAT和SOD活性,降低了超氧阴离子生成速率和花后18天MDA含量,增加了RNA和DNA含量;喷施乙烯利或氟草酮,结果相反。表明在MD处理下较高的ABA与乙烯(ACC)比值提高了膜脂过氧化保护酶(CAT、SOD)活性,增加了清除自由基(超氧阴离子)的能力,最终促进弱势粒灌浆。
4. ABA和乙烯对稻米品质的调控作用MD处理显著提高了稻米的崩解值,显著降低了垩白粒率、垩白度和消减值;SD处理导致品质变劣。乙烯释放速率与整精米率及崩解值呈负相关,与垩白米率、垩白度及消减值呈正相关;ABA/ACC与整精米率和崩解值呈正相关,与垩白度呈负相关。在花后9～13天喷施乙烯合成抑制物质硝酸钴或ABA,显著提高了整精米率、直链淀粉含量和崩解值,显著降低了稻米的垩白米率、垩白度和蛋白质含量及消减值;喷施乙烯释放促进物质乙烯利或ABA合成抑制物质氟草酮,结果则相反。表明籽粒中ABA和乙烯对水稻品质的形成起调控作用,籽粒中较高的ABA与乙烯比值有利于改善稻米品质。
Both abscisic acid (ABA) and ethylene are important hormones and play important roles in growth and development in responses to stress. It is little known, however, whether and how ABA and ethylene are involved in regulation to grain-filling in rice (Oryza sativa) and wheat (Triticum aestivum). This study investigated the roles of ABA and ethylene in grain filling in rice and wheat and their physiological mechanisms by using typical rice and wheat cultivars. The main results are as follows:
1. Regulation of ABA and ethylene to endosperm development and grain filling. The division rate and the number of endosperm cells, grain filling rate, and grain weight of superior grains in rice or wheat were greater than those of inferior grains. At the early grain filling stage, the evolution rate of ethylene and 1-aminocylopropane-1-carboxylic acid (ACC) concentrations were greater in inferior grains than in superior grains. While ABA concentrations and the ratio of ABA to ACC (ABA/ACC) were greater in the superior than in the inferior grains. The cell division and grain-filling rates were significantly and positively correlated with both ABA contents and the ratio of ABA to ACC, whereas significantly and negatively correlated with ethylene evolution rate. Application of cobalt ion (inhibitor of ethylene synthesis) or ABA at early grain filling stage (at 9-13d post-anthesis) significantly increased endosperm cell division rate and cell number, grain-filling rate and grain weight of inferior spikelets. Application of ethephon (ethylene-releasing agent) or fluridone (inhibitor of carotenoid synthesis) had the opposite effects. The results suggest that the antagonistic interactions between ABA and ethylene mediate the endosperm cell division and grain filling in rice and wheat. A higher ratio of ABA to ethylene (ACC) in rice and wheat grains is required to maintain a faster the endosperm cell division rate and grain-filling rate.
2. Responses of ABA and ethylene in rice and wheat grains to soil moisture and their relations with grain filling. Both moderate soil-drying (MD) and severe soil-drying (SD) were conducted during the grain filling period, and the treatment of well watered (WW) was taken as control. The responses of ABA and ethylene to soil drying and their relations with grain filling were studied. The results showed that, compared with WW, MD significantly increased, whereas SD significantly reduced, grain-filling rate and grain weight. ABA content in the grains was very low at the early grain filling stage, and reached a maximum when the grain-filling rate was the highest. In contrast to ABA, levels of ethylene and ACC in the grains were very high at early grain filling stage and sharply decreased during the linear period of grain growth. Both MD and SD remarkably enhanced ABA accumulation in the grains, which was enhanced more in SD grains than in MD ones. MD reduced, whereas SD remarkably increased, ethylene evolution rate and ACC content. MD increased, whereas SD reduced, the ratio of ABA to ACC. The grain-filling rates were significantly and positively correlated with both ABA contents and the ratio of ABA to ACC, whereas significantly and negatively correlated with ethylene evolution rate. Application of amino-ethoxyvinylglycine (AVG) or cobaltion (inhibitor of ethylene synthesis) or ABA at 9-13 d post-anthesis significantly reduced ACC concentrations and ethylene evolution rate of grains, and significantly enhanced the activities of sucrose synthase (SuSase), ADP glucose pyrophosphorylase (AGPase) and soluble starch synthase (SSSase), and increased grain-filling rate and grain weight. The results were reversed when ethephon (ethylene-releasing agent) or fluridone (inhibitor of ABA synthesis) was applied. The results suggested that a moderate soil-drying during the grain-filling period of rice and wheat could increase the ratio of ABA to ethylene (ACC), and therefore accelerate grain filling and increase grain weight. The mediation of ABA and ethylene to grain filling is probably through regulating key enzymes involved in sucrose-to-starch pathway, and a high ratio of ABA to ACC enhances grain filling.
3. ABA and ethylene in relation with membrane lipid peroxidation and nucleic acid content/concentration in grains at different soil moisture conditions. The MD increased activities of superoxide dismutase (SOD) and catalase (CAT). SD significantly increased those at the early treatment stage, but decreased at the late treatment stage. Among all the treatments, inferior grains had much lower activities of SOD, peroxidase (POD) and CAT than superior grains at the early grain filling stage, and the effect of soil moisture on inferior grains was much grater than on superior grains. There were no significant differences in superoxide anion (O2-) production rate and malondialdehyde (MDA) content of grains between MD and WW plants, but they were much higher in SD plants. MD significantly increased, whereas SD significantly reduced, ribonucleic acid (RNA) contents in grains. There were no significant differences in deoxyribonucleic acid (DNA) contents in grains among all the treatments. The grain-filling rates and ABA contents in grains were significantly and positively correlated with activities of SOD and CAT and RNA contents in grains, whereas significantly and negatively correlated with MDA content of grains. The ethylene evolution rate and ACC content in grains were significantly and negatively correlated with activities of SOD and CAT and RNA contents, and significantly and positively correlated with O2- production rate in grains. Application of cobaltion (inhibitor of ethylene synthesis) or ABA, at 9-13 d post-anthesis significantly enhanced the activities of SOD and CAT, increased RNA and DNA contents of grains, whereas significantly reduced O2- production rate in grains and at MDA content of grains, and the results were reversed when ethephon (ethylene-releasing agent) or fluridone (inhibitor of ABA synthesis) was applied. The results suggested that a moderate soil drying during the grain-filling period of rice and wheat could enhance the ratio of ABA to ethylene (ACC) and the activities of SOD and CAT of grains, and reduce the ethylene evolution rate and membrane lipid peroxidation in grains, and therefore accelerate the filling of inferior grains.
4. Roles of ABA and ethylene in grain quality of rice. The MD significantly increased breakdown values, and reduced chalky kernels, chalkiness and setback values. The results were reversed for the SD treatment. Ethylene evolution rates were significantly and negatively correlated with head milled rice and breakdown values, and significantly and positively correlated with chalky kernels, chalkiness and setback values. The ratio of ABA to ACC was significantly and positively correlated with head milled rice and breakdown values, and significantly and negatively correlated with chalkiness. Application of cobaltion (inhibitor of ethylene synthesis) or ABA, at 9-13 d post-anthesis significantly increased head milled rice, amylose contents and breakdown values, whereas significantly reduced chalky kernels, chalkiness, protein contents, and setback values, the results were reversed when ethephon (ethylene-releasing agent) or fluridone (inhibitor of ABA synthesis) was applied. The results indicate that Both ABA and ethylene play roles in the formation of grain quality, and that a higher ratio of ABA to ethylene (ACC) would benefit grain quality of rice.
引文
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