水稻生长对根际氧营养的响应特征及其生理机制研究
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摘要
水稻是世界上最重要的农作物,世界上一半以上的人口以水稻为主食。氧参与了稻田生态系统活动的重要过程,是水稻生长发育和生理代谢过程中主要的营养因子。水稻不但能利用通气组织和根从地上部及根际环境中吸收和转运氧以满足自身生长的需要,还能通过根系泌氧改善根际环境。稻田氧营养直接影响水稻的生长和产量的形成,具有显著的生态效应。
地表积水是导致水稻地下部(根系)缺氧胁迫的主要原因。水稻生长早期(分蘖期)需要田间淹水以防除杂草,这将导致其根际缺氧；遭受涝害时,水稻根系也会处于缺氧环境。此时,水稻根系必需在形态(如增加孔隙度)和代谢(如减少营养物质的吸收面积)上进行一定的调节,而这种调节又会影响其生长以及根际状况。控制灌水(增加土壤和空气的接触时间)是目前常见的一种水稻根际增氧途径,而利用化学物质增氧还处于试验阶段。
基于国内外研究进展,本研究拟以水稻根系形态、氮素营养和产量性状对根际氧营养的响应为突破口,兼顾水稻品种间的差异,旨在研明不同根际氧营养水稻的生物学响应及其生理机制；探寻水稻根际氧营养调控的可行途径；为水稻抗逆高产栽培提供基于氧营养调控的理论依据和技术储备。
依据水稻根系对根际氧营养的敏感程度,本课题组于2005-2006对农艺性状差异较大的近20份水稻品种进行了筛选,确定“国稻1号”和“秀水09”分别作为籼型和粳型代表品种。2006-2007,选择过氧化尿素和过氧化钙作为化学物质增氧材料,通过系列试验,验证其增氧效果并确定适宜用量。2007年通过营养液培养,确定水稻的需氧性和根际氧营养敏感时期。为验证不同增氧模式对水稻根际缺氧的调控效果,分别于2007年和2008年,设计了过氧化尿素(T1)、过氧化钙(T2)以及干湿交替灌溉(T3)等3种根际增氧模式的田间试验并以长期淹水田块为对照(CK),监测水稻地上、地下部的形态、生理特征与根际环境效应以及氮素利用效率和产量效应。获得主要结论如下：
1.过氧化尿素和过氧化钙都能与水反应,释放出氧气、提高水体的溶氧量。一次施用含20kg hm-1活性氧的两种化学物质,在不种植作物时能够使水层保持至少17d的增氧状态,且pH值升高幅度不大。水稻种植试验表明,分别在分蘖期和孕穗期追力口120kghm-2的过氧化尿素或施用364kg hm-2的过氧化钙,即共施入含40kg hm-2活性氧的两种化学物质时,水稻产量分别提高32.3%和27.5%,较其余施用量增产效果好。
2.水稻不同时期根际增氧均能提高植株的生物量,但分蘖期和孕穗期最明显。分蘖期增氧,国稻1号和秀水09生物量分别提高12.3%和44.8%,增加幅度较其他时期大。孕穗期持续低氧处理或增氧处理,水稻生物量和产量增加或降低的幅度均较其他阶段大。该阶段持续低氧处理,国稻1号和秀水09生物量分别降低14.2%和13.2%,产量分别降低28.4%和12.3%。
3.根际增氧显著促进水稻根系形态结构的优化和功能的提高,主要表现为：根系长度增加、根体积变大、根吸收面积增加和根系活力增强；增氧模式下,灌浆期水稻根系仍能保持18%以上的孔隙度,有利于根系获得较多的氧；齐穗后SOD和POD仍保持相对较强的活性,MDA(丙二醛)积累较少。
4.田间试验表明：与CK相比,国稻1号和秀水09的T1、T2和T3处理增产幅度2007年分别为：3.1%/11.5%、10.2%/14.9%和18.9%/16.4%；2008年分别为：11.56%/6.57%、8.48%/9.20%和13.56%/9.39%。在田间长期淹水条件下两种化学物质增氧处理的水稻产量接近田间干湿交替灌溉处理的水平。水稻在形态、生理及干物质积累等方面对氧营养增加的响应总体表现为：(1)前期分蘖数增加较快,有效穗增多；(2)齐穗后叶片叶绿素含量下降较慢,剑叶SOD(超氧化物歧化酶)和POD(过氧化物酶)活性较高,MDA含量较低；(3)齐穗后叶片光合作用对穗部干物质积累贡献大,结实率提高。
5.增氧模式下,水稻齐穗期叶和茎鞘中氮积累较多,增氧有利于穗部灌浆期氮的转运。齐穗后,国稻1号增氧模式的氮积累均显著高于CK,秀水09T1和T3氮积累也较CK多。收获期,两水稻品种的氮积累量均表现为T3> T1> T2> CK.增氧促进了水稻对硝态氮的吸收,且硝态氮含量品种间差异显著,其中粳稻硝态氮含量较低。叶片NRA受增氧模式的影响显著,硝酸还原酶的活性与叶片硝态氮的含量呈极显著(国稻1号)或显著(秀水09)相关。增氧模式下,水稻齐穗后仍能保持较大氮吸收能力,其机制与硝酸还原酶活性提高、氮同化加速有关。水稻的氮素偏生产力(PFPN)在两种化学物质增氧下,均有所提高,但受水稻基因型的影响,其中国稻1号的PFPN表现为T1>T2,而秀水09则相反。
6.采用Richards方程对不同增氧模式下水稻籽粒的灌浆动态进行拟合发现,增氧处理强势粒和弱势粒重均高于CK；国稻1号增氧处理强势粒的灌浆速率降低,而秀水09提高。秀水09增氧模式全穗灌浆速率较快,全穗粒重的增加主要依靠强势粒灌浆速率的提高；而国稻1号全穗灌浆速率较慢,弱势粒灌浆时间延长,全穗粒重的提高主要依靠弱势粒的贡献。
Rice (Oryza sativa L.) is the most important crop in the world, and more than half of the world population lives on it. Oxygen, as a major nutrition for rice plant, involves in the important processes of rice ecosystems and plays a great role in rice physiological metabolism and development. Rice can absorb and transport oxygen to meet the needs of its own growth through aerenchyma and roots from the aboveground and rhizosphere, and improve the rhizosphere environment through radial oxygen loss (ROL) of its root system. Oxygen nutrition in paddy directly influences rice growth and grain yield, while there are also significant ecological effects to rhizosphere environment.
Long-term flooding is the main reason causing anoxic stress to rice belowground system, especially the root systems. At early tiller stage, flooding can control weed in paddy field, but lead to hypoxia in rice rhizosphere. Oxygen deficiency of rice plant often happens while waterlodgging and flooding, especially under heavy rian fall. Characteristics of roots allowing internal aeration may conflict with those for water or nutrient acquisition, thereby, morphological and physiological adjustments are inevitable, consequently inducing adjustments to plant growing and the rhizosphere conditions. So far, alternate dry/wet irrigation, which can keep sufficient commutative area of gas between air and soil during rice growth stage, is the main method to alleviate anoxic stress in rice rhizosphere for high-yield cultivation. Besides, the studies on other oxygen-increasing patterns including aeration by peroxide application are still under experimental stage.
Accroding to the literature review of the research on oxygen nutrition in paddy at home and abroad in recent years, this study took the responses of root morphology and N uptake of rice plant to oxygen nutrition as the key topic, including the differences in the responses between rice genotypes. During2005-2006, nearly20varieties of rice, whose agronomic characteristic are quite different from each other, were compared by the sensitivity of their roots to oxygen rhizosphere in our research group, and then "Guodao1" and "Xiushui09" were selected as the representatives of indica and japonica varieties used in this reserach, respectively. In2007, a nutrition solution culture experiment was performed which aimed at determination of demand of oxygen nutrition at different stages for rice growing. During2006-2007, urea peroxide and calcium peroxide were selected as the chemical materials to improve oxygen situations in field, and their effects to rice growth and supplication were determinated by a serial of experiments. In order to monitor the morphological and physiological responses, as well as yield characteristics of rice to the three oxygen-increasing patterns in rhizosphere, two-year field trials (2007-2008) were performed. Three oxygen-increasing patterns were adopted:(1) urea peroxide application (T1), calcium peroxide application (T2) and alternate dry/wet irrigation (T3). Continuous submerging condition (no oxygen increased in rhizosphere) was taken as the control (CK). Under T1, T2and CK, a10cm depth of water was kept in the field during the whole period of rice growing.
The objectives were mainly to explore rice morphological and physiological responses to and their mechanism of different oxygen nutrition levels and oxygen regulation patterns. This study hopes to supply theoretical references and technical reserves for rice cultivation with high-yield under flooding stress. The main conclusions are as follows:
1. After reacting with water, calcium peroxide and urea peroxide can release oxygen to significantly increase dissolution oxygen content in the water. These chemicals application (equivalent to20kg ha-1active oxygen element) could keep water at high dissolution oxygen concentration for not less than17d, and the pH value was enhanced when no rice was planted in the soil. The results showed that the application amount of120kg hm-2urea peroxide or364kg hm-2at the tiller stage and booting stage respectively was the suitable doses, and the former dose increased rice yield by32.3%and the latter by27.5%.
2. Rice biomass was enhanced by oxygen-increasing rhizosphere at different growth stage, especially at the stages of tiller and booting. At the tiller stage, rice biomass increased by12.3%for'Guodaol'and44.8%for 'Xiushui09' respectively, compared with CK after20d aeration. Under hypoxia or aeration, amplitudes of variation of rice plants in yield and biomass were the largest or the smallest at booting stage than that at other stages. Keeping lower dissolution oxygen concentration in the nutrient solution from booting stage to harvest stage, rice biomass decreased by14.2%for'Guodao1' and13.2%for 'Xiushui09', grain yield by28.4%for'Guodao1'and12.3%for 'Xiushui09', respectively, comparison with the CK.
3. Results of2007-2008showed that oxygen-increasing decreased root porosity at tillering stage, while root volume and activity increased at heading stage. Aeration enhanced the activity of antioxidant enzyme, and decreased the content of MDA in rice roots.
4. The results showed that compared with the control (CK), grain yield under the treatments of T1, T2and T3increased by3.1%,10.2%.and18.9%for'guodao1',and11.5%,14.9%and16.4%for 'Xiushui09'(japonica) in2007, respectively; and by11.56%,8.48%and13.56%for the former, and6.57%,9.20%and9.39%for the later in2008, respectively. Grain yield in flooding paddy field under the two chemical materials application was near to the yield of rice in the field with alternate dry/wet irrigation. The main mechanisms underlying the yield increase under oxygen-increasing are:(1) Panicle number was larger due to the rapid development of tillers during tillering stage, and (2) Chlorophyll content of leaves decreased more slowly after heading stage, but SOD and POD activity increased while MDA concentration decreasing in leaves at harvest stage, and (3) Dry matter translocation rate was higher with greater biomass accumulation after heading.Although the three oxygen-increasing patterns had different effects on rice growth, they all could effectively alleviate the stress for root system and above-ground part.
5. Oxygen-increasing rhizosphere enhanced nitrogen use efficiency by rice plant. Under different oxygen-increasing patterns, more nitrogen was accumulated in rice leaf and stem sheath than that in roots and spikes at full heading stage, consequently may benefit N to transit to the spikes at grain-filling stage. After heading stage, nitrogen absorption by 'Guodao1'cultivar was significantly higher under oxygen-increasing treatments than under the CK, as well as, similar trends existed in 'XiushuiO9' plant under the T1or T2treatments. The nitrogen accumulation amounts of the two rice varieties at harvest stage were ranked as T3>T1>T2>CK. Nitrogen content in rice leaves was promoted by increasing-oxygen, and significantly higher nitrogen enhancement was found in the indica rice. Increasing-oxygen patterns significantly influenced NRA (nitrate reductase activity) in rice leaves. The correlations between NRA and nitrate content in rice leaves were at significant levels for 'Guodao1'(P<0.01) and 'Xiushui09'(P<0.05). Under the increasing-oxygen patterns, Oxygen-led increment in nitrate content in leaves stimulated the activity of nitrate reductase to help nitrogen assimilation of rice leaves and the latter enhance nitrogen accumulation, inducing higher nitrogen use efficiency. The PFPN(partial factor productivity from applied N) of rice under the T1and T2treatments was higher than the CK, and obvious difference existed between the genotypes. PFPN under the T1treatment for'Guodao1' was higher than that in the T2, and that under T2for 'Xiushui09' was higher than the T1.
6. Grain filling dynamic at different positions of rice panicle was investigated. The Richards equation was used to describe grain filling processes. The results showed that oxygen-increasing patterns significantly increased grain weight on lower and upper parts of panicle. The grain-filling rate of grains on upper parts of the panicle for'Guodaol'was smaller under oxygen-increasing patterns than that under the CK, but grain weight of the former was higher than the later. The grain-filling rate of 'Xiushui09' was larger than that of'Guodaol'. Time of grain-filling for'Guodaol'under oxygen-increasing patterns were prolonged, but the average rate of grain-filling was lower than CK. Amount of grain-filling for'Guodaol'was increased by oxygen-increasing resulting from the increase of grain-filling rate of grain on the lower part of panicle, but for 'Xiushui09' resulting from the increase of grain-filling rate of grain on the upper part of panicle.
地表积水是导致水稻地下部(根系)缺氧胁迫的主要原因。水稻生长早期(分蘖期)需要田间淹水以防除杂草,这将导致其根际缺氧；遭受涝害时,水稻根系也会处于缺氧环境。此时,水稻根系必需在形态(如增加孔隙度)和代谢(如减少营养物质的吸收面积)上进行一定的调节,而这种调节又会影响其生长以及根际状况。控制灌水(增加土壤和空气的接触时间)是目前常见的一种水稻根际增氧途径,而利用化学物质增氧还处于试验阶段。
基于国内外研究进展,本研究拟以水稻根系形态、氮素营养和产量性状对根际氧营养的响应为突破口,兼顾水稻品种间的差异,旨在研明不同根际氧营养水稻的生物学响应及其生理机制；探寻水稻根际氧营养调控的可行途径；为水稻抗逆高产栽培提供基于氧营养调控的理论依据和技术储备。
依据水稻根系对根际氧营养的敏感程度,本课题组于2005-2006对农艺性状差异较大的近20份水稻品种进行了筛选,确定“国稻1号”和“秀水09”分别作为籼型和粳型代表品种。2006-2007,选择过氧化尿素和过氧化钙作为化学物质增氧材料,通过系列试验,验证其增氧效果并确定适宜用量。2007年通过营养液培养,确定水稻的需氧性和根际氧营养敏感时期。为验证不同增氧模式对水稻根际缺氧的调控效果,分别于2007年和2008年,设计了过氧化尿素(T1)、过氧化钙(T2)以及干湿交替灌溉(T3)等3种根际增氧模式的田间试验并以长期淹水田块为对照(CK),监测水稻地上、地下部的形态、生理特征与根际环境效应以及氮素利用效率和产量效应。获得主要结论如下：
1.过氧化尿素和过氧化钙都能与水反应,释放出氧气、提高水体的溶氧量。一次施用含20kg hm-1活性氧的两种化学物质,在不种植作物时能够使水层保持至少17d的增氧状态,且pH值升高幅度不大。水稻种植试验表明,分别在分蘖期和孕穗期追力口120kghm-2的过氧化尿素或施用364kg hm-2的过氧化钙,即共施入含40kg hm-2活性氧的两种化学物质时,水稻产量分别提高32.3%和27.5%,较其余施用量增产效果好。
2.水稻不同时期根际增氧均能提高植株的生物量,但分蘖期和孕穗期最明显。分蘖期增氧,国稻1号和秀水09生物量分别提高12.3%和44.8%,增加幅度较其他时期大。孕穗期持续低氧处理或增氧处理,水稻生物量和产量增加或降低的幅度均较其他阶段大。该阶段持续低氧处理,国稻1号和秀水09生物量分别降低14.2%和13.2%,产量分别降低28.4%和12.3%。
3.根际增氧显著促进水稻根系形态结构的优化和功能的提高,主要表现为：根系长度增加、根体积变大、根吸收面积增加和根系活力增强；增氧模式下,灌浆期水稻根系仍能保持18%以上的孔隙度,有利于根系获得较多的氧；齐穗后SOD和POD仍保持相对较强的活性,MDA(丙二醛)积累较少。
4.田间试验表明：与CK相比,国稻1号和秀水09的T1、T2和T3处理增产幅度2007年分别为：3.1%/11.5%、10.2%/14.9%和18.9%/16.4%；2008年分别为：11.56%/6.57%、8.48%/9.20%和13.56%/9.39%。在田间长期淹水条件下两种化学物质增氧处理的水稻产量接近田间干湿交替灌溉处理的水平。水稻在形态、生理及干物质积累等方面对氧营养增加的响应总体表现为：(1)前期分蘖数增加较快,有效穗增多；(2)齐穗后叶片叶绿素含量下降较慢,剑叶SOD(超氧化物歧化酶)和POD(过氧化物酶)活性较高,MDA含量较低；(3)齐穗后叶片光合作用对穗部干物质积累贡献大,结实率提高。
5.增氧模式下,水稻齐穗期叶和茎鞘中氮积累较多,增氧有利于穗部灌浆期氮的转运。齐穗后,国稻1号增氧模式的氮积累均显著高于CK,秀水09T1和T3氮积累也较CK多。收获期,两水稻品种的氮积累量均表现为T3> T1> T2> CK.增氧促进了水稻对硝态氮的吸收,且硝态氮含量品种间差异显著,其中粳稻硝态氮含量较低。叶片NRA受增氧模式的影响显著,硝酸还原酶的活性与叶片硝态氮的含量呈极显著(国稻1号)或显著(秀水09)相关。增氧模式下,水稻齐穗后仍能保持较大氮吸收能力,其机制与硝酸还原酶活性提高、氮同化加速有关。水稻的氮素偏生产力(PFPN)在两种化学物质增氧下,均有所提高,但受水稻基因型的影响,其中国稻1号的PFPN表现为T1>T2,而秀水09则相反。
6.采用Richards方程对不同增氧模式下水稻籽粒的灌浆动态进行拟合发现,增氧处理强势粒和弱势粒重均高于CK；国稻1号增氧处理强势粒的灌浆速率降低,而秀水09提高。秀水09增氧模式全穗灌浆速率较快,全穗粒重的增加主要依靠强势粒灌浆速率的提高；而国稻1号全穗灌浆速率较慢,弱势粒灌浆时间延长,全穗粒重的提高主要依靠弱势粒的贡献。
Rice (Oryza sativa L.) is the most important crop in the world, and more than half of the world population lives on it. Oxygen, as a major nutrition for rice plant, involves in the important processes of rice ecosystems and plays a great role in rice physiological metabolism and development. Rice can absorb and transport oxygen to meet the needs of its own growth through aerenchyma and roots from the aboveground and rhizosphere, and improve the rhizosphere environment through radial oxygen loss (ROL) of its root system. Oxygen nutrition in paddy directly influences rice growth and grain yield, while there are also significant ecological effects to rhizosphere environment.
Long-term flooding is the main reason causing anoxic stress to rice belowground system, especially the root systems. At early tiller stage, flooding can control weed in paddy field, but lead to hypoxia in rice rhizosphere. Oxygen deficiency of rice plant often happens while waterlodgging and flooding, especially under heavy rian fall. Characteristics of roots allowing internal aeration may conflict with those for water or nutrient acquisition, thereby, morphological and physiological adjustments are inevitable, consequently inducing adjustments to plant growing and the rhizosphere conditions. So far, alternate dry/wet irrigation, which can keep sufficient commutative area of gas between air and soil during rice growth stage, is the main method to alleviate anoxic stress in rice rhizosphere for high-yield cultivation. Besides, the studies on other oxygen-increasing patterns including aeration by peroxide application are still under experimental stage.
Accroding to the literature review of the research on oxygen nutrition in paddy at home and abroad in recent years, this study took the responses of root morphology and N uptake of rice plant to oxygen nutrition as the key topic, including the differences in the responses between rice genotypes. During2005-2006, nearly20varieties of rice, whose agronomic characteristic are quite different from each other, were compared by the sensitivity of their roots to oxygen rhizosphere in our research group, and then "Guodao1" and "Xiushui09" were selected as the representatives of indica and japonica varieties used in this reserach, respectively. In2007, a nutrition solution culture experiment was performed which aimed at determination of demand of oxygen nutrition at different stages for rice growing. During2006-2007, urea peroxide and calcium peroxide were selected as the chemical materials to improve oxygen situations in field, and their effects to rice growth and supplication were determinated by a serial of experiments. In order to monitor the morphological and physiological responses, as well as yield characteristics of rice to the three oxygen-increasing patterns in rhizosphere, two-year field trials (2007-2008) were performed. Three oxygen-increasing patterns were adopted:(1) urea peroxide application (T1), calcium peroxide application (T2) and alternate dry/wet irrigation (T3). Continuous submerging condition (no oxygen increased in rhizosphere) was taken as the control (CK). Under T1, T2and CK, a10cm depth of water was kept in the field during the whole period of rice growing.
The objectives were mainly to explore rice morphological and physiological responses to and their mechanism of different oxygen nutrition levels and oxygen regulation patterns. This study hopes to supply theoretical references and technical reserves for rice cultivation with high-yield under flooding stress. The main conclusions are as follows:
1. After reacting with water, calcium peroxide and urea peroxide can release oxygen to significantly increase dissolution oxygen content in the water. These chemicals application (equivalent to20kg ha-1active oxygen element) could keep water at high dissolution oxygen concentration for not less than17d, and the pH value was enhanced when no rice was planted in the soil. The results showed that the application amount of120kg hm-2urea peroxide or364kg hm-2at the tiller stage and booting stage respectively was the suitable doses, and the former dose increased rice yield by32.3%and the latter by27.5%.
2. Rice biomass was enhanced by oxygen-increasing rhizosphere at different growth stage, especially at the stages of tiller and booting. At the tiller stage, rice biomass increased by12.3%for'Guodaol'and44.8%for 'Xiushui09' respectively, compared with CK after20d aeration. Under hypoxia or aeration, amplitudes of variation of rice plants in yield and biomass were the largest or the smallest at booting stage than that at other stages. Keeping lower dissolution oxygen concentration in the nutrient solution from booting stage to harvest stage, rice biomass decreased by14.2%for'Guodao1' and13.2%for 'Xiushui09', grain yield by28.4%for'Guodao1'and12.3%for 'Xiushui09', respectively, comparison with the CK.
3. Results of2007-2008showed that oxygen-increasing decreased root porosity at tillering stage, while root volume and activity increased at heading stage. Aeration enhanced the activity of antioxidant enzyme, and decreased the content of MDA in rice roots.
4. The results showed that compared with the control (CK), grain yield under the treatments of T1, T2and T3increased by3.1%,10.2%.and18.9%for'guodao1',and11.5%,14.9%and16.4%for 'Xiushui09'(japonica) in2007, respectively; and by11.56%,8.48%and13.56%for the former, and6.57%,9.20%and9.39%for the later in2008, respectively. Grain yield in flooding paddy field under the two chemical materials application was near to the yield of rice in the field with alternate dry/wet irrigation. The main mechanisms underlying the yield increase under oxygen-increasing are:(1) Panicle number was larger due to the rapid development of tillers during tillering stage, and (2) Chlorophyll content of leaves decreased more slowly after heading stage, but SOD and POD activity increased while MDA concentration decreasing in leaves at harvest stage, and (3) Dry matter translocation rate was higher with greater biomass accumulation after heading.Although the three oxygen-increasing patterns had different effects on rice growth, they all could effectively alleviate the stress for root system and above-ground part.
5. Oxygen-increasing rhizosphere enhanced nitrogen use efficiency by rice plant. Under different oxygen-increasing patterns, more nitrogen was accumulated in rice leaf and stem sheath than that in roots and spikes at full heading stage, consequently may benefit N to transit to the spikes at grain-filling stage. After heading stage, nitrogen absorption by 'Guodao1'cultivar was significantly higher under oxygen-increasing treatments than under the CK, as well as, similar trends existed in 'XiushuiO9' plant under the T1or T2treatments. The nitrogen accumulation amounts of the two rice varieties at harvest stage were ranked as T3>T1>T2>CK. Nitrogen content in rice leaves was promoted by increasing-oxygen, and significantly higher nitrogen enhancement was found in the indica rice. Increasing-oxygen patterns significantly influenced NRA (nitrate reductase activity) in rice leaves. The correlations between NRA and nitrate content in rice leaves were at significant levels for 'Guodao1'(P<0.01) and 'Xiushui09'(P<0.05). Under the increasing-oxygen patterns, Oxygen-led increment in nitrate content in leaves stimulated the activity of nitrate reductase to help nitrogen assimilation of rice leaves and the latter enhance nitrogen accumulation, inducing higher nitrogen use efficiency. The PFPN(partial factor productivity from applied N) of rice under the T1and T2treatments was higher than the CK, and obvious difference existed between the genotypes. PFPN under the T1treatment for'Guodao1' was higher than that in the T2, and that under T2for 'Xiushui09' was higher than the T1.
6. Grain filling dynamic at different positions of rice panicle was investigated. The Richards equation was used to describe grain filling processes. The results showed that oxygen-increasing patterns significantly increased grain weight on lower and upper parts of panicle. The grain-filling rate of grains on upper parts of the panicle for'Guodaol'was smaller under oxygen-increasing patterns than that under the CK, but grain weight of the former was higher than the later. The grain-filling rate of 'Xiushui09' was larger than that of'Guodaol'. Time of grain-filling for'Guodaol'under oxygen-increasing patterns were prolonged, but the average rate of grain-filling was lower than CK. Amount of grain-filling for'Guodaol'was increased by oxygen-increasing resulting from the increase of grain-filling rate of grain on the lower part of panicle, but for 'Xiushui09' resulting from the increase of grain-filling rate of grain on the upper part of panicle.
引文
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