覆盖旱种对水稻产量与品质的影响及其生理机制
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摘要
二十一世纪的全球农业面临两大挑战:一是为满足人口的增长需要不断增加粮食产量,二是在不断增加粮食产量的同时需要应对水资源的日益减少。覆膜或覆草旱种是近年来发展起来的水稻节水栽培技术,具有明显的节水效果。但对于在不同旱种方式下水稻品质形成特点的研究很少,在旱种条件下产量和稻米品质形成的生理机制尚不清楚。本研究以粳稻镇稻88、籼稻扬稻6号和杂交籼稻汕优63为材料,进行裸地旱种(NM),地膜覆盖旱种(PM)和麦秸秆覆盖旱种(SM),以常规水种(TF)为对照,分析了覆盖旱种对水稻产量和品质的影响及其生理机制。主要结果如下:
1、与TF相比,旱种水稻产量均有不同程度的下降,NM、PM和SM的减产率分别为37% ~ 49%,8.5% ~ 17%和0.3% ~ 0.7%,NM和PM的产量与TF有显著差异,SM的产量与TF差异不显著。NM的穗数、每穗粒数、结实率和千粒重均较TF显著降低,因而表现为严重减产;PM减产的原因主要在于总颖花量和千粒重的下降;SM的每穗粒数较TF显著减少,但结实率和粒重较TF显著提高;PM和NM处理的最大灌浆速率和平均灌浆速率较TF显著降低,SM处理则显著增加。表明SM处理粒重的增加与灌浆速率的增大有关,而PM和NM处理粒重降低,籽粒灌浆速率减小是重要原因。
2、在整个水稻生长期各处理的平均灌溉用水量,PM为206 mm,SM为233 mm,NM为258 mm,分别为TF(953 mm)的22%、24%和27%。所有旱种处理均较TF显著提高了灌溉水的利用效率(单位灌溉水生产的稻谷),增加的幅度PM为267% ~ 367%,SM为278% ~ 321%,NM为98% ~ 138%。表明SM在保持较高产量的同时可以大幅度地提高水分利用效率。3、SM改善了稻米的加工品质、外观品质和蒸煮品质,NM和PM则降低了稻米品质;SM提高了稻米的最高粘度和崩解值,降低了稻米的热浆粘度、最终粘度和消减值,NM和PM的结果则相反,说明SM可以改善稻米的品质。
4、与TF相比,SM提高了籽粒灌浆期的根系氧化力、光合速率和籽粒中蔗糖合成酶(SuSase),腺苷二磷酸葡萄糖焦磷酸化酶(AGPase)、淀粉合成酶(StSase)和淀粉分枝酶(SBE)活性,NM和PM则降低了上述生理参数。
5、结实期籽粒吲哚-3-乙酸(IAA)、赤霉酸(GA1+GA4)、玉米素(Z)+玉米素核苷(ZR)含量在灌浆早期处理间差异很小,在灌浆中、后期则表现为SM > TF > PM > NM,乙烯释放速率则表现为NM > PM > TF > SM。PM和SM增加了籽粒脱落酸(ABA)浓度,但与TF无显著差异,NM则显著增加了籽粒ABA含量。灌浆中后期籽粒IAA和GA1+GA4、灌浆前中期籽粒ABA及灌浆各期籽粒Z+ZR浓度与粒重、出糙率、精米率和崩解值呈显著或极显著的正相关(r = 0.71*~ 0.96**),与消减值呈显著或极显著负相关(r = -0.76* ~ -0.91**)。灌浆后期籽粒IAA和Z+ZR浓度与垩白米率和垩白度呈显著或极显著负相关(r = -0.73* ~ -0.85**)。灌浆各期籽粒乙烯释放速率与粒重、出糙率、精米率、碱解值和崩解值呈显著或极显著的负相关(r = -0.71* ~ -0.96**),与垩白米率、垩白度和消减值呈显著或极显著的正相关(r = 0.73* ~ 0.93**)。
6、SM增加了ABA与1-氨基环丙烷-1-羧酸(ACC)的比值(ABA/ACC),NM和PM则降低了ABA/ACC值,表明在NM和PM条件下乙烯的增加超过了ABA的增加。籽粒ABA含量与籽粒灌浆速率呈渐近线函数关系,乙烯释放速率与籽粒灌浆速率呈衰减的指数函数关系,ABA/ACC比值与灌浆速率则呈直线相关。在抽穗后7～10 d对TF或PM稻穗喷施乙烯合成抑制物质氨基-乙氧基乙烯基甘氨酸(AVG)或ABA,显著地降低了籽粒中乙烯的释放速率,提高了籽粒中SuSase、AGPase和可溶性StSase活性,增加了灌浆速率和粒重;喷施乙烯合成促进物质乙烯利或ABA合成抑制物质氟草酮,结果则相反。表明ABA和乙烯的相互作用调控了籽粒灌浆。
上述结果说明,覆膜旱种和覆草均可显著地提高水分利用效率。覆草旱种不仅能获得较高的产量,而且还可以改善稻米品质。覆膜旱种则降低了产量和品质。在覆草旱种条件下结实期根系活力、光合速率和籽粒中蔗糖-淀粉代谢途径关键酶活性的提高以及激素间的平衡特别是ABA与乙烯比值(ABA/ACC)的增加是其高产优质的重要生理基础。在水分短缺或易发生季节性干旱而温度不是限制水稻生长的地区,覆草旱种是较覆膜旱种更好的水稻高产优质水分高效利用的栽培技术。
Global agriculture in the 21st century faces two major challenges. Total food production needs to increase to feed a still growing world population, and this increase needs to be accomplished under increasing scarcity of water resources. Plastic film or straw mulching cultivation under non-flooded condition has been considered as a new water-saving technique in rice (Oryza sativa L.) production. This study aimed to investigate the yield performance in terms of quality and quantity, water use efficiency (WUE) and their physiological mechanism under such practices. A field experiment was conducted with three high-yielding rice cultivars, Zhendao 88 ( a japonica cultivar), Yangdao 6 (an indica cultivar) and Shanyou 63 (an indica hybrid cultivar) and four cultivation treatments imposed from transplanting to maturity: traditional flooding as control (TF), non-flooded plastic film mulching (PM), non-flooded wheat straw mulching (SM), and non-flooded no mulching (NM). Grain yield and quality, WUE and physiological parameters, such as plant hormones, activities of key enzymes involved in sucrose-starch conversion, were determined. The main results are as follows:
1. Compared with that under TF, grain yield showed some reduction under all the non-flooded cultivations, but differed largely among the treatments. The reduction was 37%-49%, 8.5%-17%, and 0.3%-0.7%, respectively, for NM, PM, and SM. The difference in grain yield was significant between NM and TF or between PM and TF, and was not significant between SM and TF. The yield reduction under NM was attributed to decreases in panicle number, spikelets per panicle, percentage of filled grains, and grain weight. The reduction in the total number of spikelets and grain weight contributed to the decrease in grain yield under PM. SM significantly reduced number of spikelets per panicle but significantly increased percentage of filled grains and grain weight. Both NM and PM significantly reduced, whereas SM significantly increased the maximum and mean grain filling rate, indicating that an increase in grain weight under SM is associated with the enhancement in grain filling rate, and decreases in grain filling rates under NM and PM result in the reduction in grain weight.
2. Water application during the growing season across the study years, on average, was 206 mm to the PM, 233 mm to the SM, and 258 mm to the NM, which was only 22%, 24%, and 27%, respectively, of that (953 mm) applied to the TF. Compared with that under the TF, irrigation water use efficiency (grain yield per unit irrigation water) was markedly increased under all the non-flooded treatments, and by 267%-367% under the PM, 278%-321% under the SM, and 98%-138% under the NM treatments. The results suggest that SM could maintain a high grain yield and markedly increase WUE.
3. SM significantly improved milling, appearance, and cooking qualities, whereas PM or NM decreased these qualities. SM also significantly increased the peak viscosity and breakdown value, and reduced hot viscosity, final viscosity, and setback value, and PM or NM had the opposite effect. Such results suggest that SM could improve the quality of rice.
4. Compared with TF, SM significantly increased root oxidation activity, photosynthetic rate, and activities of sucrose synthase (SuSase), adenosine diphosphate glucose pyrophosphorylase (AGPase), starch synthase (StSase) and starch branching enzyme (SBE) in grains during the grain filling period, whereas NM and PM significantly reduced these parameters.
5. Contents of indole-3-acetic acid (IAA), gibberellins (GA1 + GA4) and zeatin (Z) + zeatin riboside (ZR) in grains differed little among the treatments at the early grain filing stage, and showed SM > TF > PM > NM at the mid and late filling stages. Ethylene evolution rate of grains exhibited NM > PM > TF > SM. Abscisic acid (ABA) contences in grains were increased under both PM and SM, but showed no significant differences among TF, PM and SM treatments. NM significantly increased ABA in grains. Contents of IAA and GA1 + GA4 at the mid and late filling stages, ABA contents at the early and mid filling stage, and Z + ZR contents during the whole grain filling period significantly or very significantly and positively correlated with grain weight, brown rice, milled rice and breakdown value (r = 0.71* to 0.96**), and negatively correlated with setback value (r = -0.76* to -0.91**). Contents of IAA and Z + ZR at the late filling stage significantly or very significantly and negatively correlated with the percentage of chalky kernels and chalkiness (r = -0.73* to -0.85**). The ethylene evolution rates at each filling stage significantly or very significantly and negatively correlated with grain weight, brown rice, milled rice, alkali spreading value and breakdown value (r = -0.71* to -0.96**), and positively correlated with the percentage of chalky kernels, chalkiness and setback value (r = 0.73* to 0.93**).
6. SM increased the ratio of ABA to 1-aminocylopropane -1-carboxylic acid (ACC) in the grains, and both NM and PM reduced it (ABA/ACC), implying that there is a greater enhancement of ABA concentration than ethylene production in the SM treatment, whereas the increase in ethylene outweighs the increase in ABA under PM and NM treatments. The content of ABA correlated with grain filling rate with a hyperbolic curve, whereas the ethylene evolution rate correlated with grain filling rate with an exponential decay equation. The ratio of ABA to ACC significantly correlated with grain filling rate with a linear relationship. Application of amino-ethoxyvinylglycine (AVG, inhibitor of ethylene synthesis by inhibiting ACC synthase) or ABA to panicles under TF and PM at the early grain filling stage significantly increased activities of the key enzymes involved in sucrose to starch conversion in the grains, SuSase, AGPase and soluble StSase, grain filling rate and grain weight. Application of ethephon (ethylene-releasing agent) or fluridone (inhibitor of ABA synthesis) had the opposite effect. The results suggest that antagonistic interactions between ABA and ethylene may be involved in mediating the effect of non-flooded mulching cultivation on grain filling.
Collectively, both non-flooded plastic mulching cultivation (PM) and non-flooded wheat straw mulching cultivation (SM) could significantly increase WUE. SM not only maintained a high grain yield, but also improved quality of rice. Under the PM, quality of rice was decreased and grain yield differed with cultivars. Increases in photosynthetic rate, root activity, and activities of the key enzymes involved in the sucrose-starch metabolic pathway in grains, and the balance between hormones especially increase in the ratio of ABA to ethylene (ABA/ACC) under SM contributed to the high grain yield and the improvement in rice quality. SM would be a better practice than PM in areas where water is scarce but temperature is favorable to rice growth, such as the case in Southeast China.
1、与TF相比,旱种水稻产量均有不同程度的下降,NM、PM和SM的减产率分别为37% ~ 49%,8.5% ~ 17%和0.3% ~ 0.7%,NM和PM的产量与TF有显著差异,SM的产量与TF差异不显著。NM的穗数、每穗粒数、结实率和千粒重均较TF显著降低,因而表现为严重减产;PM减产的原因主要在于总颖花量和千粒重的下降;SM的每穗粒数较TF显著减少,但结实率和粒重较TF显著提高;PM和NM处理的最大灌浆速率和平均灌浆速率较TF显著降低,SM处理则显著增加。表明SM处理粒重的增加与灌浆速率的增大有关,而PM和NM处理粒重降低,籽粒灌浆速率减小是重要原因。
2、在整个水稻生长期各处理的平均灌溉用水量,PM为206 mm,SM为233 mm,NM为258 mm,分别为TF(953 mm)的22%、24%和27%。所有旱种处理均较TF显著提高了灌溉水的利用效率(单位灌溉水生产的稻谷),增加的幅度PM为267% ~ 367%,SM为278% ~ 321%,NM为98% ~ 138%。表明SM在保持较高产量的同时可以大幅度地提高水分利用效率。3、SM改善了稻米的加工品质、外观品质和蒸煮品质,NM和PM则降低了稻米品质;SM提高了稻米的最高粘度和崩解值,降低了稻米的热浆粘度、最终粘度和消减值,NM和PM的结果则相反,说明SM可以改善稻米的品质。
4、与TF相比,SM提高了籽粒灌浆期的根系氧化力、光合速率和籽粒中蔗糖合成酶(SuSase),腺苷二磷酸葡萄糖焦磷酸化酶(AGPase)、淀粉合成酶(StSase)和淀粉分枝酶(SBE)活性,NM和PM则降低了上述生理参数。
5、结实期籽粒吲哚-3-乙酸(IAA)、赤霉酸(GA1+GA4)、玉米素(Z)+玉米素核苷(ZR)含量在灌浆早期处理间差异很小,在灌浆中、后期则表现为SM > TF > PM > NM,乙烯释放速率则表现为NM > PM > TF > SM。PM和SM增加了籽粒脱落酸(ABA)浓度,但与TF无显著差异,NM则显著增加了籽粒ABA含量。灌浆中后期籽粒IAA和GA1+GA4、灌浆前中期籽粒ABA及灌浆各期籽粒Z+ZR浓度与粒重、出糙率、精米率和崩解值呈显著或极显著的正相关(r = 0.71*~ 0.96**),与消减值呈显著或极显著负相关(r = -0.76* ~ -0.91**)。灌浆后期籽粒IAA和Z+ZR浓度与垩白米率和垩白度呈显著或极显著负相关(r = -0.73* ~ -0.85**)。灌浆各期籽粒乙烯释放速率与粒重、出糙率、精米率、碱解值和崩解值呈显著或极显著的负相关(r = -0.71* ~ -0.96**),与垩白米率、垩白度和消减值呈显著或极显著的正相关(r = 0.73* ~ 0.93**)。
6、SM增加了ABA与1-氨基环丙烷-1-羧酸(ACC)的比值(ABA/ACC),NM和PM则降低了ABA/ACC值,表明在NM和PM条件下乙烯的增加超过了ABA的增加。籽粒ABA含量与籽粒灌浆速率呈渐近线函数关系,乙烯释放速率与籽粒灌浆速率呈衰减的指数函数关系,ABA/ACC比值与灌浆速率则呈直线相关。在抽穗后7～10 d对TF或PM稻穗喷施乙烯合成抑制物质氨基-乙氧基乙烯基甘氨酸(AVG)或ABA,显著地降低了籽粒中乙烯的释放速率,提高了籽粒中SuSase、AGPase和可溶性StSase活性,增加了灌浆速率和粒重;喷施乙烯合成促进物质乙烯利或ABA合成抑制物质氟草酮,结果则相反。表明ABA和乙烯的相互作用调控了籽粒灌浆。
上述结果说明,覆膜旱种和覆草均可显著地提高水分利用效率。覆草旱种不仅能获得较高的产量,而且还可以改善稻米品质。覆膜旱种则降低了产量和品质。在覆草旱种条件下结实期根系活力、光合速率和籽粒中蔗糖-淀粉代谢途径关键酶活性的提高以及激素间的平衡特别是ABA与乙烯比值(ABA/ACC)的增加是其高产优质的重要生理基础。在水分短缺或易发生季节性干旱而温度不是限制水稻生长的地区,覆草旱种是较覆膜旱种更好的水稻高产优质水分高效利用的栽培技术。
Global agriculture in the 21st century faces two major challenges. Total food production needs to increase to feed a still growing world population, and this increase needs to be accomplished under increasing scarcity of water resources. Plastic film or straw mulching cultivation under non-flooded condition has been considered as a new water-saving technique in rice (Oryza sativa L.) production. This study aimed to investigate the yield performance in terms of quality and quantity, water use efficiency (WUE) and their physiological mechanism under such practices. A field experiment was conducted with three high-yielding rice cultivars, Zhendao 88 ( a japonica cultivar), Yangdao 6 (an indica cultivar) and Shanyou 63 (an indica hybrid cultivar) and four cultivation treatments imposed from transplanting to maturity: traditional flooding as control (TF), non-flooded plastic film mulching (PM), non-flooded wheat straw mulching (SM), and non-flooded no mulching (NM). Grain yield and quality, WUE and physiological parameters, such as plant hormones, activities of key enzymes involved in sucrose-starch conversion, were determined. The main results are as follows:
1. Compared with that under TF, grain yield showed some reduction under all the non-flooded cultivations, but differed largely among the treatments. The reduction was 37%-49%, 8.5%-17%, and 0.3%-0.7%, respectively, for NM, PM, and SM. The difference in grain yield was significant between NM and TF or between PM and TF, and was not significant between SM and TF. The yield reduction under NM was attributed to decreases in panicle number, spikelets per panicle, percentage of filled grains, and grain weight. The reduction in the total number of spikelets and grain weight contributed to the decrease in grain yield under PM. SM significantly reduced number of spikelets per panicle but significantly increased percentage of filled grains and grain weight. Both NM and PM significantly reduced, whereas SM significantly increased the maximum and mean grain filling rate, indicating that an increase in grain weight under SM is associated with the enhancement in grain filling rate, and decreases in grain filling rates under NM and PM result in the reduction in grain weight.
2. Water application during the growing season across the study years, on average, was 206 mm to the PM, 233 mm to the SM, and 258 mm to the NM, which was only 22%, 24%, and 27%, respectively, of that (953 mm) applied to the TF. Compared with that under the TF, irrigation water use efficiency (grain yield per unit irrigation water) was markedly increased under all the non-flooded treatments, and by 267%-367% under the PM, 278%-321% under the SM, and 98%-138% under the NM treatments. The results suggest that SM could maintain a high grain yield and markedly increase WUE.
3. SM significantly improved milling, appearance, and cooking qualities, whereas PM or NM decreased these qualities. SM also significantly increased the peak viscosity and breakdown value, and reduced hot viscosity, final viscosity, and setback value, and PM or NM had the opposite effect. Such results suggest that SM could improve the quality of rice.
4. Compared with TF, SM significantly increased root oxidation activity, photosynthetic rate, and activities of sucrose synthase (SuSase), adenosine diphosphate glucose pyrophosphorylase (AGPase), starch synthase (StSase) and starch branching enzyme (SBE) in grains during the grain filling period, whereas NM and PM significantly reduced these parameters.
5. Contents of indole-3-acetic acid (IAA), gibberellins (GA1 + GA4) and zeatin (Z) + zeatin riboside (ZR) in grains differed little among the treatments at the early grain filing stage, and showed SM > TF > PM > NM at the mid and late filling stages. Ethylene evolution rate of grains exhibited NM > PM > TF > SM. Abscisic acid (ABA) contences in grains were increased under both PM and SM, but showed no significant differences among TF, PM and SM treatments. NM significantly increased ABA in grains. Contents of IAA and GA1 + GA4 at the mid and late filling stages, ABA contents at the early and mid filling stage, and Z + ZR contents during the whole grain filling period significantly or very significantly and positively correlated with grain weight, brown rice, milled rice and breakdown value (r = 0.71* to 0.96**), and negatively correlated with setback value (r = -0.76* to -0.91**). Contents of IAA and Z + ZR at the late filling stage significantly or very significantly and negatively correlated with the percentage of chalky kernels and chalkiness (r = -0.73* to -0.85**). The ethylene evolution rates at each filling stage significantly or very significantly and negatively correlated with grain weight, brown rice, milled rice, alkali spreading value and breakdown value (r = -0.71* to -0.96**), and positively correlated with the percentage of chalky kernels, chalkiness and setback value (r = 0.73* to 0.93**).
6. SM increased the ratio of ABA to 1-aminocylopropane -1-carboxylic acid (ACC) in the grains, and both NM and PM reduced it (ABA/ACC), implying that there is a greater enhancement of ABA concentration than ethylene production in the SM treatment, whereas the increase in ethylene outweighs the increase in ABA under PM and NM treatments. The content of ABA correlated with grain filling rate with a hyperbolic curve, whereas the ethylene evolution rate correlated with grain filling rate with an exponential decay equation. The ratio of ABA to ACC significantly correlated with grain filling rate with a linear relationship. Application of amino-ethoxyvinylglycine (AVG, inhibitor of ethylene synthesis by inhibiting ACC synthase) or ABA to panicles under TF and PM at the early grain filling stage significantly increased activities of the key enzymes involved in sucrose to starch conversion in the grains, SuSase, AGPase and soluble StSase, grain filling rate and grain weight. Application of ethephon (ethylene-releasing agent) or fluridone (inhibitor of ABA synthesis) had the opposite effect. The results suggest that antagonistic interactions between ABA and ethylene may be involved in mediating the effect of non-flooded mulching cultivation on grain filling.
Collectively, both non-flooded plastic mulching cultivation (PM) and non-flooded wheat straw mulching cultivation (SM) could significantly increase WUE. SM not only maintained a high grain yield, but also improved quality of rice. Under the PM, quality of rice was decreased and grain yield differed with cultivars. Increases in photosynthetic rate, root activity, and activities of the key enzymes involved in the sucrose-starch metabolic pathway in grains, and the balance between hormones especially increase in the ratio of ABA to ethylene (ABA/ACC) under SM contributed to the high grain yield and the improvement in rice quality. SM would be a better practice than PM in areas where water is scarce but temperature is favorable to rice growth, such as the case in Southeast China.
引文
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