超级稻产量形成生理与栽培调控的研究
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摘要
追求高产、更高产是水稻生产发展的永恒主题。研究超级稻产量形成生理及其栽培调控对提高我国水稻综合生产能力,保障粮食安全供给具有重要的意义。本研究以代表性超级稻品种为材料,对超级稻的冠层结构、叶片光合特性和根系形态生理特征与产量的关系、超级稻花后强、弱势粒淀粉合成相关酶活性和激素含量变化及其与籽粒灌浆的关系、超级稻抽穗前茎中储存的非结构性碳水化合物与库强的关系、水分和氮肥对超级稻产量及品质的影响及其生理机制、低温和弱光胁迫对超级稻产量和品质的影响及其生理机制等进行了研究。主要结果如下：
1.超级稻的冠层结构、叶片光合特性及根系形态生理特征与产量形成的关系
以4个超级稻品种两优培九和Ⅱ优084(杂交籼稻)、淮稻9号和武粳15(粳稻)为材料,2个高产品种汕优63(杂交籼稻)和扬辐粳8号(粳稻)为对照进行研究。4个超级稻品种的平均每穗粒数和产量较两个对照品种分别高出57.19%和16.13%,但超级稻的结实率较对照品种低15.26个百分点。超级稻品种每穗粒数的增加主要表现在二次枝梗数目和二次枝梗总粒数的增加,结实率低则主要表现在二次枝梗上籽粒的结实率较低。超级稻品种具有较高的分蘖成穗率和透光率、较大的叶面积指数和粒叶比、较小的顶部3片叶的叶基角、齐穗期与成熟期较高的生物产量。超级稻品种在生育前、中期叶片中叶绿素a含量、叶绿素b含量、总叶绿素含量、类胡萝卜素含量、光合速率、蒸腾速率、气孔导度和根系中单位干重根系活力、每株根系活力、总根系吸收面积、活跃吸收面积和比表面积均高于对照品种,而在生育后期以上性状指标下降速率要大于各自对照品种,直至齐穗后20d前以上性状指标均小于各自对照品种。说明超级稻强大的产量库容与其生育前、中期较强的叶片光合能力和较好的根系生理性状密切有关,生育后期叶片光合能力和根系生理活性下降快导致了其结实率的下降,从而限制了其产量潜力的发挥。
2.超级稻花后强、弱势粒淀粉合成相关酶活性和激素含量变化及其与籽粒灌浆的关系
研究了以上4个超级稻品种和2个高产品种的花后强、弱势粒淀粉合成相关酶活性和激素含量变化及其与籽粒灌浆的关系。结果表明,4个超级稻品种强势粒的最大灌浆速率、到达最大灌浆速率的时间、平均灌浆速率和糙米重与各自对照品种相比差异较小,超级稻品种弱势粒的灌浆速率和糙米重表现为显著低于各自对照品种。灌浆期强、弱势粒的蔗糖合酶(SuSase)、腺苷二磷酸葡萄糖焦磷酸化酶(AGPase)、淀粉合酶(StSase)和淀粉分支酶(SBE)活性变化和玉米素+玉米素核苷(Z+ZR)、吲哚-3-乙酸(IAA)和脱落酸(ABA)含量变化均呈单峰值曲线。弱势粒的SuSase、AGPase、StSase和SBE的峰值活性和平均活性及其Z+ZR和IAA的含量峰值浓度和平均含量均低于强势粒。弱势粒的ABA含量峰值和平均含量显著高于强势粒,超级稻品种高于对照品种。籽粒灌浆速率与SuSase、AGPase和StSase活性及Z+ZR和IAA含量均呈显著或极显著正相关,与SBE活性及ABA含量的相关不显著。说明超级稻品种弱势粒中较低的SuSase、AGPase和StSase活性及较低的Z+ZR和IAA含量是其灌浆速率小、粒重轻的一个重要生理原因。
3.超级稻抽穗前茎鞘中储存的非结构性碳水化合物与库强的关系
两个超级稻品种两优培九和淮稻9号、两个新株型品系IR65600-127-6-2-3(NPT1)和IR65598-112-2(NPT2)及两个常规水稻品种扬稻6号和镇稻88种植于大田,观察了抽穗前茎鞘中储存的非结构性碳水化合物与库强的关系。与常规品种相比较,超级稻和新株型稻因每穗颖花数较多而表现出较大的产量潜力,但其弱势粒较低的籽粒灌浆速率和较小的籽粒重量导致了较低的结实率。超级稻和新株型稻库容小(胚乳细胞数少)和灌浆初期库活性低[蔗糖合酶(SuSase)口腺苷二磷酸葡萄糖焦磷酸化酶(AGPase)舌性低]是弱势粒灌浆速率小和粒重低的重要原因。抽穗前茎鞘中非结构性碳水化合物(NSC)的累积量或抽穗期糖花比(NSC/颖花)与弱势粒的库强(胚乳细胞数目、SuSase和AGPase活性)、籽粒灌浆速率和籽粒重量呈显著的正相关。在颖花分化期施用氮肥可以显著增加抽穗期糖花比、超级稻弱势粒的库强、籽粒灌浆速率和籽粒重量,而在穗分化始期施氮或者在穗分化始期和颖花分化期均施氮的结果则相反。说明抽穗前茎鞘中储存的非结构性碳水化合物与水稻籽粒灌浆期库强密切相关,在颖花分化期适当施氮是增加抽穗前非结构性碳水化合物储存的技术措施。
4.结实期干湿交替灌溉对超级稻结实率和粒重的影响
以2个超级稻品种两优培九和淮稻9号为材料,自抽穗至成熟设置种灌溉方式处理：轻干-湿交替灌溉(WMD),重干-湿交替灌溉(WSD)和常规灌溉(CI,保持水层)。结果表明,与CI相比,WMD处理显著增加了两个超级稻品种的结实率和粒重,而WSD处理则降低了结实率和粒重。与CI相比,WMD处理显著提高了灌浆期剑叶净光合速率、膜质过氧化酶活性和根系氧化力、根系吸收表面积、根系活跃吸收表面积、比表面积和根系激素中玉米素+玉米素核苷(Z+ZR)及吲哚-3-乙酸(IAA)含量和根冠比,而WSD处理的结果则相反。说明在灌浆期进行轻干-湿交替灌溉,可以提高超级稻的结实率和粒重。
5.氮肥对超级稻产量形成的调控作用
以两个超级稻品种两优培九和淮稻9号为材料,以两个高产品种汕优63和扬辐粳8号为对照,设置四种氮肥水平,即：全生育期不施氮肥(TO)、全生育期施氮120kg hm-2(T1)、全生育期施氮240kg hm-2(T2)和全生育期施氮360kg hm-2(T3)。在施氮量0～360kg hm-2条件下,两个超级稻品种的产量随着施氮量的增加而增加,灌浆期剑叶光合速率(Pn)和硝酸还原酶活性(NR)、根系氧化活力和玉米素+玉米素核苷(Z+ZR)含量、地上部干物重、根干重和根冠比均随着施氮量的增加而增加。而两个对照品种的产量在施氮量0～240kg hm-2的条件下,产量随施氮量的增加而增加,灌浆期剑叶Pn和NR活性、根系氧化活力和Z+ZR含量、地上部干物重、根干重和根冠比均随着施氮量的增加而增加。当施氮量继续增加至360kg hm-2时,产量则明显下降,灌浆期剑叶Pn和NR活性、根系氧化活力和Z+ZR含量、地上部干物重、根干重和根冠比则显著下降。说明通过氮肥调控,可以促进超级稻根-冠生长,进而获得高产。
6.抽穗-灌浆期低温与弱光胁迫对超级稻籽粒充实的影响
以两个超级稻品种两优培九和淮稻9号为材料,以两个高产品种汕优63和扬辐粳8号为对照,设置两种低温胁迫：即自抽穗至抽穗后10d20℃低温处理(T1)；抽穗后11d至20d20℃低温处理(T2)及两种弱光胁迫处理：自抽穗至抽穗后10d40%遮光处理(透光率为60%)(T3)；抽穗后11d至20d40%遮光处理(透光率为60%)(T4)。以自然温度、自然光照为对照(CK)。低温、弱光胁迫显著降低了4个水稻品种的产量和结实率,两个超级稻品种的下降幅度要大于各自对照品种。剑叶光合速率(Pn)、过氧化氢酶(CAT)活性、超氧化物歧化酶(SOD)活性和根干重、根系氧化活力在灌浆期均呈下降趋势,低温、弱光胁迫显著降低了以上指标,两个超级稻品种的下降幅度均要大于各自对照品种。剑叶的丙二醛(MDA)含量和Pn变化趋势相反。过氧化物酶(POD)活性从抽穗-抽穗后20d变化趋势和CAT变化趋势一致,但到花后30d时,又呈现出上升趋势,低温、弱光胁迫显著增加了POD的活性,两个超级稻表现更加明显。根系中玉米素+玉米素核苷(Z+ZR)含量在灌浆期呈现出先上升后下降的趋势,低温、弱光胁迫显著降低了Z+ZR含量,两个超级稻表现更加明显。其中,20℃的低温胁迫对水稻产量和生理机制的影响要大于40%遮光(透光率为60%)的弱光胁迫的影响,抽穗至抽穗后10d的低温和弱光胁迫对水稻产量和生理机制的影响要大于抽穗后11d-20d的低温和弱光胁迫的影响。
7.超级稻稻米品质及环境对其的影响
以超级稻品种两优培九和Ⅱ优084(杂交籼稻)、淮稻9号和武粳15(粳稻)为材料,以2个高产品种汕优63和扬辐粳8号为对照,大田种植。测定了稻米品质。结果表明,超级稻品种的糙米率、精米率、整精米率和稻米中胶稠度和淀粉谱的最高黏度、热浆黏度、崩解值和蛋白组分中清蛋白、谷蛋白和米粉中赖氨酸、蛋氨酸、苏氨酸、异亮氨酸、苯丙氨酸、精氨酸、甘氨酸、丙氨酸、天冬氨酸、谷氨酸的含量均高于或显著高于各自对照品种,而超级稻品种的垩白粒率、垩白度和稻米的直链淀粉含量、蛋白质含量和淀粉谱的消碱值、回复值和蛋白组分的球蛋白、醇溶蛋白和米粉中缬氨酸、亮氨酸、组氨酸、络氨酸、脯氨酸含量均低于或显著低于常规高产品种。说明了超级稻品种具有良好的稻米品质。与常规灌溉(CI,保持水层)相比,轻干-湿交替灌溉(WMD)显著改善了稻米品质,重干-湿交替灌溉(WSD)则降低了稻米品质。在施氮量较低时(<240kg hm-2),增施氮肥有利于提高稻米品质,在施氮量为360kg hm-2时,稻米品质下降。低温和弱光处理对稻米品质有不利影响,对超级稻米质的影响大于对常规高产品种米质的影响。
综上所述,本研究从冠层结构、叶片光合生产能力和根系形态生理特性等方面阐明了超级稻产量形成生理,从库强与籽粒灌浆关系等方面阐明了超级稻结实率低的生理原因,明确了提高超级稻产量和品质的肥水调控技术,获得了超级稻产量形成生理的新认识,对超级稻高产、优质、高效栽培具有重要指导意义。
Striving for high-yielding and super-high-yielding is the perpetual topic in the development of the rice production. Studies on yield formation physiology and cultivation regulation approaches in super rice would have great importance in both improving the overall production capacity of rice in China and ensuring food safety supplies. Using typical super rice cultivars as materials, this study investigated changes in canopy structure, leaf photosynthetic characteristics, morphological and physiological traits of root and their relationships with shoot growth, post-anthesis changes in activities of the enzymes related to starch synthesis and contents of hormones in superior and inferior spikelets and their relations with grain filling, pre-anthesis non-structural carbohydrate reserve in the stem in relations with the sink strength of inferior spikelets during grain filling, effect of water and nitrogen management on yield and quality and its physiological mechanism, effect of low temperature and light stress on yield and quality in super rice and its physiological mechanism. The main results are as follows:
1. Relationship of canopy structure, leaf photosynthetic characteristics and root physiological traits with grain yield formation in super rice
Four super rice cultivars, Liangyoupeijiu and Ⅱ you084(indica hybrids), Huaidao9and Wujing15(japonica), and two elite high-yielding check cultivars, Shanyou63(indica hybrids) and Yangfujing8(japonica), were investigated. The average spikelets per panicle and grain yield of the four super rice cultivars were higher57.19%and16.13%, respectively, than those of two elite check cultivars, but the seed-setting rate of the four super rice cultivars were lower15.26%than that of the two elite check cultivars. The increase in spikelets per panicle of the four super rice cultivars were attributed to the increase of secondary branch number and secondary branch grain number, and lower seed-setting rate was mainly resulted from lower seed setting rate on the secondary branch. Super rice had higher percentage of productive tillers and light transmission rate, leaf area index and grain-leaf area ratio, smaller basic angle of top three leaves, higher percentage of matter transportation and biomass during full heading stage and maturity stage. The contents of chlorophyll a and b, total chlorophyll (a+b) and carotenoid, photosynthetic rate, transpiration rate, stomata conductance of leaves, and specific root oxidation activity, root oxidation activity per plant, total root absorption area per plant, root active absorption area, specific surface area were higher for super rice than for elite check rice at earlier and the mid-growth stages, while these traits decreased more quickly in super rice than in check rice at the latter growth stage, and these traits were lower in super rice than in check rice from20days after full heading. The results suggest that stronger leaf photosynthetic capacity and root activity during the early and mid-growth periods contribute to greater yield sink capacity of super rice. While the rapid decline of leaf photosynthetic capacity and root physiological activity at the late growth stage could result in poor grain filling, and consequently, limit yield potential of super nce.
2. Post-anthesis changes in activities of the enzymes related to starch synthesis and contents of hormones in superior and inferior spikelets and their relations with grain filling of super rice
Post-anthesis changes in activities of the enzymes related to starch synthesis and contents of honnones in superior and inferior spikelets and their relation with grain filling of the four super rice cultivars and two high-yielding cultivars were investigated. The results showed that the maximum grain filling rate, the time reaching a maximum grain-filling rate, mean grain filling rate and brown rice weight for superior spikelets showed a rather small difference between super rice and elite check rice, but those for inferior spikelets were significantly lower in super rice than in check rice cultivars. Activities of sucrose synthase (SuSase), adenosine diphosphoglucose pyrophosphorylase (AGPase), starch synthase (StSase), and starch branching enzyme (SBE) and contents of zeatin+zeatin riboside (Z+ZR), indole-3-acetic acid (IAA), abscisic acid (ABA) in superior and inferior spikelets exhibited single peak value curve during grain filling stage. The peak value and mean activity of SuSase, AGPase, StSase, SBE and the content of Z+ZR and IAA were lower in inferior than in superior spikelets. The peak and mean contents of ABA were significantly greater in inferior than in superior spikelets and greater in the super rice than in the check rice. The grain filling rate was positively significantly or very significantly correlated with the activity of SuSase, AGPase and StSase and contents of Z+ZR and IAA, but not significantly correlated with activities of SBE and ABA content. The results suggested that the low activity of SuSase, AGPase and StSase and low contents of Z+ZR and IAA may be an important physiological reason for the small grain filling rate and low grain weight of inferior spikelets in super nee.
3. Pre-anthesis non-structural carbohydrate reserve in the stem in relation with the sink strength of inferior spikelets during grain filling of rice
Two super rice varieties, Liangyoupeijiu and Huaidao9, two New Plant Type rice lines, IR65600-127-6-2-3(NPT1) and IR65598-112-2(NPT2) were compared with two conventional rice varieties under field-grown conditions, and the relationship of pre-anthesis non-structural carbohydrate reserve in the stem with the sink strength was investigated. Compared with conventional rice varieties, super rice and NPT rice showed a greater yield capacity as a result of larger spikelets per panicle, however, slow grain filling rate and low grain weight in inferior spikelets resulted in low seed-setting rate. The low grain filling rate and small grain weight of inferior spikelets are mainly attributed to a poor sink strength as a result of small sink size (small number of endosperm cells) and low sink activity such as low activities of sucrose synthase (SuSase) and adenosine diphosphoglucose pyrophosphorylase (AGPase) during early grain filling stage for super rice and NPT rice. The amount of NSC in the stem and NSC per spikelet at the heading time are significantly and positively correlated with sink strength (number of endosperm cells and activities of SuSase and AGPase), grain filling rate, and grain weight of inferior spikelets. Nitrogen application at the spikelet differentiation stage significantly increased, whereas N application at the panicle initiation or at both panicle initiation and spikelet differentiation stage, significantly reduced, NSC per spikelet at the heading time, sink strength, grain filling rate, and grain weight of inferior spikelet in super rice. The results suggest that pre-anthesis NSC reserves in the stem are closely associated with the sink strength during grain filling of rice, and N application at the spikelet differentiation stage would be a good technical measure to increase pre-anthesis NSC reserves.
4. Effects of alternate wetting and drying irrigation during the grain filling stage on seed-setting rate and grain weight of super rice
Two super rice cultivars, Liangyoupeijiu and Huaidao9were used for the study. Three irrigation regimes, alternate wetting and moderate soil drying (WMD), alternate wetting and severe soil drying (WSD), and conventional irrigation (CI, continuous flooding), were imposed from heading to maturity. Compared with the CI, seed setting rate and grain weight of super rice were significantly increased in the WMD, whereas significantly reduced in the WSD. Compared with the CI, net photosynthetic rate, membrane lipid peroxidation of flag leaf and root oxidation activity, root absorption surface area, root active absorption area, root specific surface area, zeatin+zeatin riboside (Z+ZR) and indole-3-acetic acid (IAA) in root hormone and root/shoot value in the WMD were significantly increased during grain filling stage, whereas the results were reverse in the WSD. The results suggest that the WMD imposed during grain filling could improve seed-setting rate and grain weight of super rice.
5. Regulatory role of nitrogen management on yield formation of super rice
Two super rice cultivars, Liangyoupeijiu and Huaidao9, two high-yielding elite check cultivars, Shanyou63(indica hybrid) and Yangfujing8(japonica), were used for the study. Four nitrogen (N) treatments, no N application (TO) and the N tate at120kg hm-2(T1), at240kg hm-2(T2), and at360kg hm-2(T3), were conducted. In the condition of N rate from0to360kg hm-2, grain yield of two super rice cultivars was increased with the increase in N application. Similarly, the net photosynthetic rate (Pn) and nitrate reductase (NR) activity of the flag leaf, root oxidation activity and zeatin+zeatin riboside (Z+ZR) content, shoot dry weight, root dry weight and root/shoot value during the grain filling period were increased with the increase in N application. At the condition of N rate from0to240kg hm-2, grain yield of two check cultivars was increased with the increase in N application, Pn and NR activity of the flag leaf, root oxidation activity and Z+ZR content, shoot dry weight, root dry weight and root/shoot value during grain filling were also increased with the increase in N application. When the N rate was at360kg hm-2, the grain yield of two check cultivars were decreased significantly, so were the Pn and NR activity of flag leaf, root oxidation activity and Z+ZR content, shoot dry weight, root dry weight and root/shoot value. The results suggest that root-shoot growth and grain yield of super rice could be improved through nitrogen management.
6. Effects of low temperature and weak light stress during heading and grain filling on grain plumpness of super rice
Two super rice cultivars, Liangyoupeijiu and Huaidao9, two high-yielding and elite check cultivars, Shanyou63and Yangfujing8, were used for study. Two low temperature stress treatments,20℃treatment from heading to10days after heading (T1),20℃treatment from11days to20days after heading (T2). Two weak light stress treatments,40%shading treatment (light transmission rate of60%) from heading to10days after heading (T3),40%shading treatment (light transmission rate of60%) from11days to20days after heading (T4). Natural temperature and natural light were taken as the control (CK). Low temperature and weak light stress significantly decreased grain yield and seed-setting rate of four rice cultivars, the decline of the two super rice cultivars was greater than their respective control cultivars. Net photosynthetic rate (Pn), catalase (CAT) activity, superoxide dismutase (SOD) activity of the flag leaf and root dry weight, root oxidation activity decreased during grain filling stage, low temperature and weak light stress significantly decreased the above indexes, the decline of the two super rice cultivars was greater than their respective control cultivars. The change of malondialdehyde (MDA) content in the flag leaf was opposite to the change of Pn. The change of peroxidase (POD) activity was consistent to the change of CAT from heading to20days after heading, while POD activity showed an inctrase trend at30days after heading. The Low temperature and weak light stress significantly increased POD activity, and increased more for the two super rice cultivars. Zeatin+zeatin riboside (Z+ZR) content in root was increased first and then decreased during grain filling. The low temperature and weak light stress significantly decreased Z+ZR content. The effect of20℃low temperature stress on grain yield and physiological mechanism in rice was greater than the effect of40%shading treatment, the effect of low temperature and weak light stress on grain yield and physiological mechanism in rice from heading to10days after heading was greater than the effect of low temperature and weak light stress from11days to20days after heading.
7. Rice quality of super rice and the influence of environment on it
Four super rice cultivars, Liangyoupeijiu and Ⅱ you084(indica hybrids), Huaidao9and Wujing15(japonica), and two elite check cultivars, Shanyou63(indica hybrids) and Yangfujing8(japonica) were field-grown, and rice quality were investigated. The results showed that brown rice rate, milled rice rate, head milled rice rate, gel consistency, peak viscosity, hot viscosity, breakdown, the content of albumin, glutelin, lysine, methionine, threonine, isoleucine, phenylalanine, arginine, glycine, alanine, asparagine, glutamate in super rice cultivars were higher or significantly higher than their respective control cultivars, while chalky grain, chalkiness, amylase content, protein content, setback, consistence the content of globulin, prolamin, valine, leucine, histidine, tyrosine, proline were lower or significantly lower than their respective control cultivars. The results suggest that super rice cultivars have good rice quality. Compared with the conventional irrigation (CI, continuous flooding), alternate wetting and moderate soil drying (WMD) significantly improved rice quality, while alternate wetting and severe soil drying (WSD) significantly reduced rice quality. Increase in N application was beneficial to improve rice quality when N rate was low (<240kg hm-2), while rice quality declined when N rate was at360kg hm-2. Low temperature and weak light stress had adverse effect on rice quality and the effect on rice quality of super rice was greater than elite high-yielding cultivars.
Collectively, this study has illustrated yield formation physiology in super rice from the aspects of canopy structure, leaf photosynthetic characteristics and root morphology and physiology, and provided information to understand the physiological reasons of low seed-setting rate in super rice from the relationships between sink strength and grain filling, and established fertilizer and water management for improving grain yield and rice quality of super rice. This study has obtained novel insights in yield formation physiology of super rice, which would have great significance in guiding cultivation for high yield, good quality, and high efficiency in super rice.
1.超级稻的冠层结构、叶片光合特性及根系形态生理特征与产量形成的关系
以4个超级稻品种两优培九和Ⅱ优084(杂交籼稻)、淮稻9号和武粳15(粳稻)为材料,2个高产品种汕优63(杂交籼稻)和扬辐粳8号(粳稻)为对照进行研究。4个超级稻品种的平均每穗粒数和产量较两个对照品种分别高出57.19%和16.13%,但超级稻的结实率较对照品种低15.26个百分点。超级稻品种每穗粒数的增加主要表现在二次枝梗数目和二次枝梗总粒数的增加,结实率低则主要表现在二次枝梗上籽粒的结实率较低。超级稻品种具有较高的分蘖成穗率和透光率、较大的叶面积指数和粒叶比、较小的顶部3片叶的叶基角、齐穗期与成熟期较高的生物产量。超级稻品种在生育前、中期叶片中叶绿素a含量、叶绿素b含量、总叶绿素含量、类胡萝卜素含量、光合速率、蒸腾速率、气孔导度和根系中单位干重根系活力、每株根系活力、总根系吸收面积、活跃吸收面积和比表面积均高于对照品种,而在生育后期以上性状指标下降速率要大于各自对照品种,直至齐穗后20d前以上性状指标均小于各自对照品种。说明超级稻强大的产量库容与其生育前、中期较强的叶片光合能力和较好的根系生理性状密切有关,生育后期叶片光合能力和根系生理活性下降快导致了其结实率的下降,从而限制了其产量潜力的发挥。
2.超级稻花后强、弱势粒淀粉合成相关酶活性和激素含量变化及其与籽粒灌浆的关系
研究了以上4个超级稻品种和2个高产品种的花后强、弱势粒淀粉合成相关酶活性和激素含量变化及其与籽粒灌浆的关系。结果表明,4个超级稻品种强势粒的最大灌浆速率、到达最大灌浆速率的时间、平均灌浆速率和糙米重与各自对照品种相比差异较小,超级稻品种弱势粒的灌浆速率和糙米重表现为显著低于各自对照品种。灌浆期强、弱势粒的蔗糖合酶(SuSase)、腺苷二磷酸葡萄糖焦磷酸化酶(AGPase)、淀粉合酶(StSase)和淀粉分支酶(SBE)活性变化和玉米素+玉米素核苷(Z+ZR)、吲哚-3-乙酸(IAA)和脱落酸(ABA)含量变化均呈单峰值曲线。弱势粒的SuSase、AGPase、StSase和SBE的峰值活性和平均活性及其Z+ZR和IAA的含量峰值浓度和平均含量均低于强势粒。弱势粒的ABA含量峰值和平均含量显著高于强势粒,超级稻品种高于对照品种。籽粒灌浆速率与SuSase、AGPase和StSase活性及Z+ZR和IAA含量均呈显著或极显著正相关,与SBE活性及ABA含量的相关不显著。说明超级稻品种弱势粒中较低的SuSase、AGPase和StSase活性及较低的Z+ZR和IAA含量是其灌浆速率小、粒重轻的一个重要生理原因。
3.超级稻抽穗前茎鞘中储存的非结构性碳水化合物与库强的关系
两个超级稻品种两优培九和淮稻9号、两个新株型品系IR65600-127-6-2-3(NPT1)和IR65598-112-2(NPT2)及两个常规水稻品种扬稻6号和镇稻88种植于大田,观察了抽穗前茎鞘中储存的非结构性碳水化合物与库强的关系。与常规品种相比较,超级稻和新株型稻因每穗颖花数较多而表现出较大的产量潜力,但其弱势粒较低的籽粒灌浆速率和较小的籽粒重量导致了较低的结实率。超级稻和新株型稻库容小(胚乳细胞数少)和灌浆初期库活性低[蔗糖合酶(SuSase)口腺苷二磷酸葡萄糖焦磷酸化酶(AGPase)舌性低]是弱势粒灌浆速率小和粒重低的重要原因。抽穗前茎鞘中非结构性碳水化合物(NSC)的累积量或抽穗期糖花比(NSC/颖花)与弱势粒的库强(胚乳细胞数目、SuSase和AGPase活性)、籽粒灌浆速率和籽粒重量呈显著的正相关。在颖花分化期施用氮肥可以显著增加抽穗期糖花比、超级稻弱势粒的库强、籽粒灌浆速率和籽粒重量,而在穗分化始期施氮或者在穗分化始期和颖花分化期均施氮的结果则相反。说明抽穗前茎鞘中储存的非结构性碳水化合物与水稻籽粒灌浆期库强密切相关,在颖花分化期适当施氮是增加抽穗前非结构性碳水化合物储存的技术措施。
4.结实期干湿交替灌溉对超级稻结实率和粒重的影响
以2个超级稻品种两优培九和淮稻9号为材料,自抽穗至成熟设置种灌溉方式处理：轻干-湿交替灌溉(WMD),重干-湿交替灌溉(WSD)和常规灌溉(CI,保持水层)。结果表明,与CI相比,WMD处理显著增加了两个超级稻品种的结实率和粒重,而WSD处理则降低了结实率和粒重。与CI相比,WMD处理显著提高了灌浆期剑叶净光合速率、膜质过氧化酶活性和根系氧化力、根系吸收表面积、根系活跃吸收表面积、比表面积和根系激素中玉米素+玉米素核苷(Z+ZR)及吲哚-3-乙酸(IAA)含量和根冠比,而WSD处理的结果则相反。说明在灌浆期进行轻干-湿交替灌溉,可以提高超级稻的结实率和粒重。
5.氮肥对超级稻产量形成的调控作用
以两个超级稻品种两优培九和淮稻9号为材料,以两个高产品种汕优63和扬辐粳8号为对照,设置四种氮肥水平,即：全生育期不施氮肥(TO)、全生育期施氮120kg hm-2(T1)、全生育期施氮240kg hm-2(T2)和全生育期施氮360kg hm-2(T3)。在施氮量0～360kg hm-2条件下,两个超级稻品种的产量随着施氮量的增加而增加,灌浆期剑叶光合速率(Pn)和硝酸还原酶活性(NR)、根系氧化活力和玉米素+玉米素核苷(Z+ZR)含量、地上部干物重、根干重和根冠比均随着施氮量的增加而增加。而两个对照品种的产量在施氮量0～240kg hm-2的条件下,产量随施氮量的增加而增加,灌浆期剑叶Pn和NR活性、根系氧化活力和Z+ZR含量、地上部干物重、根干重和根冠比均随着施氮量的增加而增加。当施氮量继续增加至360kg hm-2时,产量则明显下降,灌浆期剑叶Pn和NR活性、根系氧化活力和Z+ZR含量、地上部干物重、根干重和根冠比则显著下降。说明通过氮肥调控,可以促进超级稻根-冠生长,进而获得高产。
6.抽穗-灌浆期低温与弱光胁迫对超级稻籽粒充实的影响
以两个超级稻品种两优培九和淮稻9号为材料,以两个高产品种汕优63和扬辐粳8号为对照,设置两种低温胁迫：即自抽穗至抽穗后10d20℃低温处理(T1)；抽穗后11d至20d20℃低温处理(T2)及两种弱光胁迫处理：自抽穗至抽穗后10d40%遮光处理(透光率为60%)(T3)；抽穗后11d至20d40%遮光处理(透光率为60%)(T4)。以自然温度、自然光照为对照(CK)。低温、弱光胁迫显著降低了4个水稻品种的产量和结实率,两个超级稻品种的下降幅度要大于各自对照品种。剑叶光合速率(Pn)、过氧化氢酶(CAT)活性、超氧化物歧化酶(SOD)活性和根干重、根系氧化活力在灌浆期均呈下降趋势,低温、弱光胁迫显著降低了以上指标,两个超级稻品种的下降幅度均要大于各自对照品种。剑叶的丙二醛(MDA)含量和Pn变化趋势相反。过氧化物酶(POD)活性从抽穗-抽穗后20d变化趋势和CAT变化趋势一致,但到花后30d时,又呈现出上升趋势,低温、弱光胁迫显著增加了POD的活性,两个超级稻表现更加明显。根系中玉米素+玉米素核苷(Z+ZR)含量在灌浆期呈现出先上升后下降的趋势,低温、弱光胁迫显著降低了Z+ZR含量,两个超级稻表现更加明显。其中,20℃的低温胁迫对水稻产量和生理机制的影响要大于40%遮光(透光率为60%)的弱光胁迫的影响,抽穗至抽穗后10d的低温和弱光胁迫对水稻产量和生理机制的影响要大于抽穗后11d-20d的低温和弱光胁迫的影响。
7.超级稻稻米品质及环境对其的影响
以超级稻品种两优培九和Ⅱ优084(杂交籼稻)、淮稻9号和武粳15(粳稻)为材料,以2个高产品种汕优63和扬辐粳8号为对照,大田种植。测定了稻米品质。结果表明,超级稻品种的糙米率、精米率、整精米率和稻米中胶稠度和淀粉谱的最高黏度、热浆黏度、崩解值和蛋白组分中清蛋白、谷蛋白和米粉中赖氨酸、蛋氨酸、苏氨酸、异亮氨酸、苯丙氨酸、精氨酸、甘氨酸、丙氨酸、天冬氨酸、谷氨酸的含量均高于或显著高于各自对照品种,而超级稻品种的垩白粒率、垩白度和稻米的直链淀粉含量、蛋白质含量和淀粉谱的消碱值、回复值和蛋白组分的球蛋白、醇溶蛋白和米粉中缬氨酸、亮氨酸、组氨酸、络氨酸、脯氨酸含量均低于或显著低于常规高产品种。说明了超级稻品种具有良好的稻米品质。与常规灌溉(CI,保持水层)相比,轻干-湿交替灌溉(WMD)显著改善了稻米品质,重干-湿交替灌溉(WSD)则降低了稻米品质。在施氮量较低时(<240kg hm-2),增施氮肥有利于提高稻米品质,在施氮量为360kg hm-2时,稻米品质下降。低温和弱光处理对稻米品质有不利影响,对超级稻米质的影响大于对常规高产品种米质的影响。
综上所述,本研究从冠层结构、叶片光合生产能力和根系形态生理特性等方面阐明了超级稻产量形成生理,从库强与籽粒灌浆关系等方面阐明了超级稻结实率低的生理原因,明确了提高超级稻产量和品质的肥水调控技术,获得了超级稻产量形成生理的新认识,对超级稻高产、优质、高效栽培具有重要指导意义。
Striving for high-yielding and super-high-yielding is the perpetual topic in the development of the rice production. Studies on yield formation physiology and cultivation regulation approaches in super rice would have great importance in both improving the overall production capacity of rice in China and ensuring food safety supplies. Using typical super rice cultivars as materials, this study investigated changes in canopy structure, leaf photosynthetic characteristics, morphological and physiological traits of root and their relationships with shoot growth, post-anthesis changes in activities of the enzymes related to starch synthesis and contents of hormones in superior and inferior spikelets and their relations with grain filling, pre-anthesis non-structural carbohydrate reserve in the stem in relations with the sink strength of inferior spikelets during grain filling, effect of water and nitrogen management on yield and quality and its physiological mechanism, effect of low temperature and light stress on yield and quality in super rice and its physiological mechanism. The main results are as follows:
1. Relationship of canopy structure, leaf photosynthetic characteristics and root physiological traits with grain yield formation in super rice
Four super rice cultivars, Liangyoupeijiu and Ⅱ you084(indica hybrids), Huaidao9and Wujing15(japonica), and two elite high-yielding check cultivars, Shanyou63(indica hybrids) and Yangfujing8(japonica), were investigated. The average spikelets per panicle and grain yield of the four super rice cultivars were higher57.19%and16.13%, respectively, than those of two elite check cultivars, but the seed-setting rate of the four super rice cultivars were lower15.26%than that of the two elite check cultivars. The increase in spikelets per panicle of the four super rice cultivars were attributed to the increase of secondary branch number and secondary branch grain number, and lower seed-setting rate was mainly resulted from lower seed setting rate on the secondary branch. Super rice had higher percentage of productive tillers and light transmission rate, leaf area index and grain-leaf area ratio, smaller basic angle of top three leaves, higher percentage of matter transportation and biomass during full heading stage and maturity stage. The contents of chlorophyll a and b, total chlorophyll (a+b) and carotenoid, photosynthetic rate, transpiration rate, stomata conductance of leaves, and specific root oxidation activity, root oxidation activity per plant, total root absorption area per plant, root active absorption area, specific surface area were higher for super rice than for elite check rice at earlier and the mid-growth stages, while these traits decreased more quickly in super rice than in check rice at the latter growth stage, and these traits were lower in super rice than in check rice from20days after full heading. The results suggest that stronger leaf photosynthetic capacity and root activity during the early and mid-growth periods contribute to greater yield sink capacity of super rice. While the rapid decline of leaf photosynthetic capacity and root physiological activity at the late growth stage could result in poor grain filling, and consequently, limit yield potential of super nce.
2. Post-anthesis changes in activities of the enzymes related to starch synthesis and contents of hormones in superior and inferior spikelets and their relations with grain filling of super rice
Post-anthesis changes in activities of the enzymes related to starch synthesis and contents of honnones in superior and inferior spikelets and their relation with grain filling of the four super rice cultivars and two high-yielding cultivars were investigated. The results showed that the maximum grain filling rate, the time reaching a maximum grain-filling rate, mean grain filling rate and brown rice weight for superior spikelets showed a rather small difference between super rice and elite check rice, but those for inferior spikelets were significantly lower in super rice than in check rice cultivars. Activities of sucrose synthase (SuSase), adenosine diphosphoglucose pyrophosphorylase (AGPase), starch synthase (StSase), and starch branching enzyme (SBE) and contents of zeatin+zeatin riboside (Z+ZR), indole-3-acetic acid (IAA), abscisic acid (ABA) in superior and inferior spikelets exhibited single peak value curve during grain filling stage. The peak value and mean activity of SuSase, AGPase, StSase, SBE and the content of Z+ZR and IAA were lower in inferior than in superior spikelets. The peak and mean contents of ABA were significantly greater in inferior than in superior spikelets and greater in the super rice than in the check rice. The grain filling rate was positively significantly or very significantly correlated with the activity of SuSase, AGPase and StSase and contents of Z+ZR and IAA, but not significantly correlated with activities of SBE and ABA content. The results suggested that the low activity of SuSase, AGPase and StSase and low contents of Z+ZR and IAA may be an important physiological reason for the small grain filling rate and low grain weight of inferior spikelets in super nee.
3. Pre-anthesis non-structural carbohydrate reserve in the stem in relation with the sink strength of inferior spikelets during grain filling of rice
Two super rice varieties, Liangyoupeijiu and Huaidao9, two New Plant Type rice lines, IR65600-127-6-2-3(NPT1) and IR65598-112-2(NPT2) were compared with two conventional rice varieties under field-grown conditions, and the relationship of pre-anthesis non-structural carbohydrate reserve in the stem with the sink strength was investigated. Compared with conventional rice varieties, super rice and NPT rice showed a greater yield capacity as a result of larger spikelets per panicle, however, slow grain filling rate and low grain weight in inferior spikelets resulted in low seed-setting rate. The low grain filling rate and small grain weight of inferior spikelets are mainly attributed to a poor sink strength as a result of small sink size (small number of endosperm cells) and low sink activity such as low activities of sucrose synthase (SuSase) and adenosine diphosphoglucose pyrophosphorylase (AGPase) during early grain filling stage for super rice and NPT rice. The amount of NSC in the stem and NSC per spikelet at the heading time are significantly and positively correlated with sink strength (number of endosperm cells and activities of SuSase and AGPase), grain filling rate, and grain weight of inferior spikelets. Nitrogen application at the spikelet differentiation stage significantly increased, whereas N application at the panicle initiation or at both panicle initiation and spikelet differentiation stage, significantly reduced, NSC per spikelet at the heading time, sink strength, grain filling rate, and grain weight of inferior spikelet in super rice. The results suggest that pre-anthesis NSC reserves in the stem are closely associated with the sink strength during grain filling of rice, and N application at the spikelet differentiation stage would be a good technical measure to increase pre-anthesis NSC reserves.
4. Effects of alternate wetting and drying irrigation during the grain filling stage on seed-setting rate and grain weight of super rice
Two super rice cultivars, Liangyoupeijiu and Huaidao9were used for the study. Three irrigation regimes, alternate wetting and moderate soil drying (WMD), alternate wetting and severe soil drying (WSD), and conventional irrigation (CI, continuous flooding), were imposed from heading to maturity. Compared with the CI, seed setting rate and grain weight of super rice were significantly increased in the WMD, whereas significantly reduced in the WSD. Compared with the CI, net photosynthetic rate, membrane lipid peroxidation of flag leaf and root oxidation activity, root absorption surface area, root active absorption area, root specific surface area, zeatin+zeatin riboside (Z+ZR) and indole-3-acetic acid (IAA) in root hormone and root/shoot value in the WMD were significantly increased during grain filling stage, whereas the results were reverse in the WSD. The results suggest that the WMD imposed during grain filling could improve seed-setting rate and grain weight of super rice.
5. Regulatory role of nitrogen management on yield formation of super rice
Two super rice cultivars, Liangyoupeijiu and Huaidao9, two high-yielding elite check cultivars, Shanyou63(indica hybrid) and Yangfujing8(japonica), were used for the study. Four nitrogen (N) treatments, no N application (TO) and the N tate at120kg hm-2(T1), at240kg hm-2(T2), and at360kg hm-2(T3), were conducted. In the condition of N rate from0to360kg hm-2, grain yield of two super rice cultivars was increased with the increase in N application. Similarly, the net photosynthetic rate (Pn) and nitrate reductase (NR) activity of the flag leaf, root oxidation activity and zeatin+zeatin riboside (Z+ZR) content, shoot dry weight, root dry weight and root/shoot value during the grain filling period were increased with the increase in N application. At the condition of N rate from0to240kg hm-2, grain yield of two check cultivars was increased with the increase in N application, Pn and NR activity of the flag leaf, root oxidation activity and Z+ZR content, shoot dry weight, root dry weight and root/shoot value during grain filling were also increased with the increase in N application. When the N rate was at360kg hm-2, the grain yield of two check cultivars were decreased significantly, so were the Pn and NR activity of flag leaf, root oxidation activity and Z+ZR content, shoot dry weight, root dry weight and root/shoot value. The results suggest that root-shoot growth and grain yield of super rice could be improved through nitrogen management.
6. Effects of low temperature and weak light stress during heading and grain filling on grain plumpness of super rice
Two super rice cultivars, Liangyoupeijiu and Huaidao9, two high-yielding and elite check cultivars, Shanyou63and Yangfujing8, were used for study. Two low temperature stress treatments,20℃treatment from heading to10days after heading (T1),20℃treatment from11days to20days after heading (T2). Two weak light stress treatments,40%shading treatment (light transmission rate of60%) from heading to10days after heading (T3),40%shading treatment (light transmission rate of60%) from11days to20days after heading (T4). Natural temperature and natural light were taken as the control (CK). Low temperature and weak light stress significantly decreased grain yield and seed-setting rate of four rice cultivars, the decline of the two super rice cultivars was greater than their respective control cultivars. Net photosynthetic rate (Pn), catalase (CAT) activity, superoxide dismutase (SOD) activity of the flag leaf and root dry weight, root oxidation activity decreased during grain filling stage, low temperature and weak light stress significantly decreased the above indexes, the decline of the two super rice cultivars was greater than their respective control cultivars. The change of malondialdehyde (MDA) content in the flag leaf was opposite to the change of Pn. The change of peroxidase (POD) activity was consistent to the change of CAT from heading to20days after heading, while POD activity showed an inctrase trend at30days after heading. The Low temperature and weak light stress significantly increased POD activity, and increased more for the two super rice cultivars. Zeatin+zeatin riboside (Z+ZR) content in root was increased first and then decreased during grain filling. The low temperature and weak light stress significantly decreased Z+ZR content. The effect of20℃low temperature stress on grain yield and physiological mechanism in rice was greater than the effect of40%shading treatment, the effect of low temperature and weak light stress on grain yield and physiological mechanism in rice from heading to10days after heading was greater than the effect of low temperature and weak light stress from11days to20days after heading.
7. Rice quality of super rice and the influence of environment on it
Four super rice cultivars, Liangyoupeijiu and Ⅱ you084(indica hybrids), Huaidao9and Wujing15(japonica), and two elite check cultivars, Shanyou63(indica hybrids) and Yangfujing8(japonica) were field-grown, and rice quality were investigated. The results showed that brown rice rate, milled rice rate, head milled rice rate, gel consistency, peak viscosity, hot viscosity, breakdown, the content of albumin, glutelin, lysine, methionine, threonine, isoleucine, phenylalanine, arginine, glycine, alanine, asparagine, glutamate in super rice cultivars were higher or significantly higher than their respective control cultivars, while chalky grain, chalkiness, amylase content, protein content, setback, consistence the content of globulin, prolamin, valine, leucine, histidine, tyrosine, proline were lower or significantly lower than their respective control cultivars. The results suggest that super rice cultivars have good rice quality. Compared with the conventional irrigation (CI, continuous flooding), alternate wetting and moderate soil drying (WMD) significantly improved rice quality, while alternate wetting and severe soil drying (WSD) significantly reduced rice quality. Increase in N application was beneficial to improve rice quality when N rate was low (<240kg hm-2), while rice quality declined when N rate was at360kg hm-2. Low temperature and weak light stress had adverse effect on rice quality and the effect on rice quality of super rice was greater than elite high-yielding cultivars.
Collectively, this study has illustrated yield formation physiology in super rice from the aspects of canopy structure, leaf photosynthetic characteristics and root morphology and physiology, and provided information to understand the physiological reasons of low seed-setting rate in super rice from the relationships between sink strength and grain filling, and established fertilizer and water management for improving grain yield and rice quality of super rice. This study has obtained novel insights in yield formation physiology of super rice, which would have great significance in guiding cultivation for high yield, good quality, and high efficiency in super rice.
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