磷对东北粳稻耐冷性及产量和品质的影响
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摘要
低温冷害是我国东北地区水稻减产的最主要气候因素之一，在各时期低温影响中，尤以移栽期低温造成的生育延迟和孕穗期低温引起的结实率下降对生产危害更大。目前，生产上普遍采用减氮增磷技术来防御低温冷害，但有关磷素提高水稻耐冷机理的研究报道较少。本研究以强耐冷水稻品种吉粳81和弱耐冷品种长白9为供试材料，通过人工气候模拟低温胁迫，设计移栽期延迟性冷害和孕穗期障碍性冷害，并设计相应的磷肥施用梯度，研究低温胁迫下磷肥施用水平对东北粳稻耐冷生理特性及产量和品质的影响。主要结果如下：
1.移栽期低温影响吉粳81和长白9秧苗的生长，导致最高分蘖时间延迟，分蘖数减少，新叶出叶速度减缓。低温下床土适当增施磷肥，两品种最高分蘖数分别提高19.7%和27.4%，新叶出叶速度分别提高8.55%和10.7%，最高分蘖发生时间分别缩短至2.7天和3.9天，成熟期秆长伸缩率缩短9.15%和10.40%。强耐冷性品种吉粳81赤枯率的变化幅度较小，弱耐冷性品种长白9的赤枯率变化明显，耐低温级数从3级上升到1级。表明床土适当增施磷肥可提高东北粳稻抵御延迟性冷害的能力。
2.孕穗期低温使吉粳81和长白9结实率、成熟期株高以及穗抽出度下降。低温下基肥中适当增施磷肥，两品种成熟期株高分别提高12.4%和15.4%，穗抽出度分别增加0.77倍和6.63倍，减轻了低温伤害的影响。结实率吉强耐冷性品种粳81的变化范围为72.3-88.0%，耐冷级数由3级上升到1级，而弱耐冷性品种长白9变化范围为52.3-78.3%，耐冷级数由5级上升到3级。说明基肥适当增施磷肥可提高东北粳稻抵御障碍性冷害的能力。
3.低温胁迫下，两品种移栽期净光合速率(Pn)分别下降79.5%和69.8%，孕穗期分别下降12.8%和19.4%。荧光动力学分析显示，随着施磷量的提高，两品种的Fv/Fm降幅不断减少，其中强耐冷品种吉粳81qP升高，NPQ下降，在施磷量120kg/hm~2时分别达到最大值0.70和最小值0.37，弱耐冷性品种长白9，qP升高，NPQ也升高，分别在施磷量120kg/hm~2和160kg/hm~2时达到最大值0.68和0.46，表明施磷提高了PSII反应中心对激发能的捕获能力和光合电子的传递能力，也增强了弱耐冷品种光保护能力，通过较高的热耗散减轻过剩激发能对反映中心的伤害。
4.低温胁迫下，适当增施磷肥可使水稻电解质外渗率和MDA下降，提高水稻叶片可溶性糖、脯氨酸含量、过氧化物酶(POD)、过氧化氢酶(CAT)、超氧化物歧化酶(SOD)的活性。相比强耐冷品种吉粳81，增磷使长白9叶片抗氧化酶活性提高幅度更大，其中POD活性增加38.2%、CAT活性增加39.7%，SOD活性增加98.9%，表明适当增施磷肥可明显降低叶片质膜透性，提高抗氧化酶活性，从而有利于提高东北粳稻抵御障碍性冷害的能力。
5.受移栽期低温影响，吉粳81和长白9二次枝梗数、穗长、每穴穗数、每穗实粒数和千粒重下降，其对产量构成因素影响大小依次为每穗实粒数＞每穴穗数＞千粒重，在床土中增施磷肥可增加二次枝梗数数、穗长、每穴穗数、每穗实粒数和千粒重。受孕穗期低温的影响，也可使吉两品种二次枝梗数、穗长、每穴穗数、每穗实粒数和千粒重下降。其对产量构成因素的影响依次为每穗实粒数＞千粒重＞每穴穗数。基肥中增施磷肥可提高两品种穗长、二次枝梗数数、每穴穗数、每穗实粒数和千粒重，且对弱耐冷品种影响尤为明显。
6.低温下，施磷可提高两品种糙米率、精米率、整精米率、垩白度、脂肪酸含量和直链淀粉含量，降低垩白米率和蛋白质含量。相关分析表明，常温下各品质指标与施磷量相关性均未达到显著水平。而低温下吉粳81整精米率与施磷量呈极显著正相关，长白9糙米率、精米率和整精米率呈显著正相关。两品种的蛋白质含量与施磷量呈负相关，脂肪酸含量呈极显著正相关，其他各品质指标相关不显著。
7.综合考虑产量及各因素状况，强耐冷品种和弱耐冷品种床土最适施磷范围分别为61.69g/m~2－71.18g/m~2和60.35g/m~2－68.02g/m~2，可消减延迟性冷害对水稻生育进程的影响；基肥最适施磷范围分别为103.65kg/hm~2－120.44kg/hm~2和91.85kg/hm~2－114.73kg/hm~2，可提高东北粳稻抵御障碍性冷害的能力。尤其在低温易发生地区生产上应尽量接近施磷范围上限，可在一定程度上减轻低温对产量的影响。
Low temperature stress frequent occurred in Jilin province which has resulted insubstantial drop in rice production for seven times in our history. At present, famers generallyuse less nitrogen but more phosphate and spray water and phosphate in production to defenselow temperature, but the inner mechanism concerning the process has rarely been reported.Therefore, the study takes cold tolerance rice varieties Japonica81and Changbai9as studyobjects; simulates low temperature condition in rice transplanting and booting periods anddesigns corresponding phosphate fertilizer gradient through artificial climate facilities;analyzes the physiology and yield of the two rice varieties under different degrees of lowtemperature and phosphate ranges, determines optimal phosphate range in different growthperiods under low temperature. The main results are as follows:
I．Low temperature in transplanting period will affect the growth of Japonica81andChangbai9seedlings and lead to the reduction of the maximum number of tillers, delay of themaximum tiller time and slower speed of new leaves growth. However, if adding appropriateamount of phosphate fertilizer in the bed soil under low temperature, the maximum number oftillers improve19.7%and27.4%, new leaves improve8.55%and10.7%, the maximumtillering time reduce2.7d and3.9d, expansion rate of stalk in maturity shorten9.15and10.40%. Seedling mortality of Jijing81change less than Changbai9, low temperatureresistance level of Changbai9is from3level to1level. The result show that soil increasedphosphate fertilizer can improve cold tolerance ability in northeast japonica rice.
II．Low temperature in booting stage results in decline in seed setting rate, mature plantheight and headings of Jijing81and Changbai9. However, adding phosphate fertilizer in thebasal, mature plant heigh of two varieties increase12.4%and15.4%, headings increase0.77times and6.63times, seed setting rate of Jijing81is range from72.3~88.0%, low temperatureresistance level is from3level to1level, but seed setting rate of Changbai9is range from52.3~78.3%, low temperature resistance level of is from5level to3level, The result showthat increased phosphate fertilizer can improve cold tolerance ability in northeast japonicarice.
Ⅲ. Net photosynthetic rate of two varieties decrease79.5%and69.8%in transplantstage under low temperature, and in booting stage they decrease12.8%and19.4%.Fluorescence dynamic analysis shows that Fv/Fm decrease by adding phosphate fertilizer. ForJijing81, qP in crease and NPQ decrease, when phosphate fertilizer application is120kg/hm~2,qP reach maximum(0.70)and NPQ reach minimum(0.37). the reasults show that cold resistantvarieties of Jijing81can use higher NPQ to dissipate. For Changbai9,qP and NPQ increase,when phosphate fertilizer application is120kg/hm~2, it reach maximum. The reasults showthat Changbai9can use higher qP to reduce reflect the center of damag.but NPQ is increase,so increaseing phosphate fertilizer application can increase the radiation heat dissipation ofenergy, reduce the damage of low temperature stress, improve the northeast japonica rice coldtolerance ability.
Ⅳ. Under the low temperature, increasing phosphate fertilizer application，membranepermeability and MDA decreased，but soluble sugar content，proline, peroxidase(POD),catalase(CAT) and superoxide dismutase (SOD) increased. Compared with Jijing81, PODactivity in Changbai9is increased higher when applying more P. The activity of POD, CATand SOD are increased38.2%,39.7%and98.9%accordingly. The results indicate thatappropriate increase the amount of fertilizer P will enhance membrane permeability of leavesand the activity of POD, so that increase the cold disaster resistance of Japonica rice innortheast China.
Ⅴ. The low temperature in transplanting period causes output decrease of Japonica81and Changbai9in many aspects such as secondary branch number, ear length, panicles perhole, grains per panicle and grain weight, among which the influential order follows as grainsper panicle> panicles per hole>grain weight. However, after adding Phosphate fertilizer inthe bed soil, the above mentioned factors improve greatly for the two varieties. The lowtemperature in booting stage causes decrease of Japonica81and Changbai9in many aspectssuch as secondary branch number, ear length, panicles per hole, grains per panicle and grainweight. But different from the influential factors orders in transplanting period, the order inbooting stage follows as grains per panicle> grain weight> panicles per hole. However, afteradding phosphate fertilizer in the bed soil in booting stage, the above mentioned factors improve greatly for both varieties and phosphates exert more effects on non-chilling tolerantvarieties.
Ⅵ. Under low temperature condition, phosphate can increase the brown rice rate, milledrice rate, chalkiness, the fatty acid content and the straight-chain starch content of the two ricevarieties. It can also lower chalky grain rate and protein content. According to related analysis,the quality indicators are not closely related to phosphate levels under normal temperaturecondition. However, under low temperature condition, the high milled rice rate of Japonica81reflects its significant positive correlation with phosphate level. As to Changbai9, its brownrice rate, milled rice rate and complete milled rice rate are all in high positive correlation tophosphate level. In contrast, the protein content of both rice varieties are negatively related tophosphate amount but the fatty acid content is significantly positively related to Phosphatefertilizer amount. Other quality indicators are not significantly related to phosphate.
Ⅶ. Considering the yield and other influential factors, the optimal phosphates in bed soilranges for Japonica81and Changbai9are respectively61.69g/m~2~71.18g/m~2and60.35g/m~2~68.02g/m~2. It can reduce effect of the growing process in rice under lowtemperature stress. The optimal basic phosphate manure ranges respectively from103.65kg/hm~2~120.44kg/hm~2to91.85kg/hm~2~114.73kg/hm~2. It can improve cold toleranceability.
1.移栽期低温影响吉粳81和长白9秧苗的生长，导致最高分蘖时间延迟，分蘖数减少，新叶出叶速度减缓。低温下床土适当增施磷肥，两品种最高分蘖数分别提高19.7%和27.4%，新叶出叶速度分别提高8.55%和10.7%，最高分蘖发生时间分别缩短至2.7天和3.9天，成熟期秆长伸缩率缩短9.15%和10.40%。强耐冷性品种吉粳81赤枯率的变化幅度较小，弱耐冷性品种长白9的赤枯率变化明显，耐低温级数从3级上升到1级。表明床土适当增施磷肥可提高东北粳稻抵御延迟性冷害的能力。
2.孕穗期低温使吉粳81和长白9结实率、成熟期株高以及穗抽出度下降。低温下基肥中适当增施磷肥，两品种成熟期株高分别提高12.4%和15.4%，穗抽出度分别增加0.77倍和6.63倍，减轻了低温伤害的影响。结实率吉强耐冷性品种粳81的变化范围为72.3-88.0%，耐冷级数由3级上升到1级，而弱耐冷性品种长白9变化范围为52.3-78.3%，耐冷级数由5级上升到3级。说明基肥适当增施磷肥可提高东北粳稻抵御障碍性冷害的能力。
3.低温胁迫下，两品种移栽期净光合速率(Pn)分别下降79.5%和69.8%，孕穗期分别下降12.8%和19.4%。荧光动力学分析显示，随着施磷量的提高，两品种的Fv/Fm降幅不断减少，其中强耐冷品种吉粳81qP升高，NPQ下降，在施磷量120kg/hm~2时分别达到最大值0.70和最小值0.37，弱耐冷性品种长白9，qP升高，NPQ也升高，分别在施磷量120kg/hm~2和160kg/hm~2时达到最大值0.68和0.46，表明施磷提高了PSII反应中心对激发能的捕获能力和光合电子的传递能力，也增强了弱耐冷品种光保护能力，通过较高的热耗散减轻过剩激发能对反映中心的伤害。
4.低温胁迫下，适当增施磷肥可使水稻电解质外渗率和MDA下降，提高水稻叶片可溶性糖、脯氨酸含量、过氧化物酶(POD)、过氧化氢酶(CAT)、超氧化物歧化酶(SOD)的活性。相比强耐冷品种吉粳81，增磷使长白9叶片抗氧化酶活性提高幅度更大，其中POD活性增加38.2%、CAT活性增加39.7%，SOD活性增加98.9%，表明适当增施磷肥可明显降低叶片质膜透性，提高抗氧化酶活性，从而有利于提高东北粳稻抵御障碍性冷害的能力。
5.受移栽期低温影响，吉粳81和长白9二次枝梗数、穗长、每穴穗数、每穗实粒数和千粒重下降，其对产量构成因素影响大小依次为每穗实粒数＞每穴穗数＞千粒重，在床土中增施磷肥可增加二次枝梗数数、穗长、每穴穗数、每穗实粒数和千粒重。受孕穗期低温的影响，也可使吉两品种二次枝梗数、穗长、每穴穗数、每穗实粒数和千粒重下降。其对产量构成因素的影响依次为每穗实粒数＞千粒重＞每穴穗数。基肥中增施磷肥可提高两品种穗长、二次枝梗数数、每穴穗数、每穗实粒数和千粒重，且对弱耐冷品种影响尤为明显。
6.低温下，施磷可提高两品种糙米率、精米率、整精米率、垩白度、脂肪酸含量和直链淀粉含量，降低垩白米率和蛋白质含量。相关分析表明，常温下各品质指标与施磷量相关性均未达到显著水平。而低温下吉粳81整精米率与施磷量呈极显著正相关，长白9糙米率、精米率和整精米率呈显著正相关。两品种的蛋白质含量与施磷量呈负相关，脂肪酸含量呈极显著正相关，其他各品质指标相关不显著。
7.综合考虑产量及各因素状况，强耐冷品种和弱耐冷品种床土最适施磷范围分别为61.69g/m~2－71.18g/m~2和60.35g/m~2－68.02g/m~2，可消减延迟性冷害对水稻生育进程的影响；基肥最适施磷范围分别为103.65kg/hm~2－120.44kg/hm~2和91.85kg/hm~2－114.73kg/hm~2，可提高东北粳稻抵御障碍性冷害的能力。尤其在低温易发生地区生产上应尽量接近施磷范围上限，可在一定程度上减轻低温对产量的影响。
Low temperature stress frequent occurred in Jilin province which has resulted insubstantial drop in rice production for seven times in our history. At present, famers generallyuse less nitrogen but more phosphate and spray water and phosphate in production to defenselow temperature, but the inner mechanism concerning the process has rarely been reported.Therefore, the study takes cold tolerance rice varieties Japonica81and Changbai9as studyobjects; simulates low temperature condition in rice transplanting and booting periods anddesigns corresponding phosphate fertilizer gradient through artificial climate facilities;analyzes the physiology and yield of the two rice varieties under different degrees of lowtemperature and phosphate ranges, determines optimal phosphate range in different growthperiods under low temperature. The main results are as follows:
I．Low temperature in transplanting period will affect the growth of Japonica81andChangbai9seedlings and lead to the reduction of the maximum number of tillers, delay of themaximum tiller time and slower speed of new leaves growth. However, if adding appropriateamount of phosphate fertilizer in the bed soil under low temperature, the maximum number oftillers improve19.7%and27.4%, new leaves improve8.55%and10.7%, the maximumtillering time reduce2.7d and3.9d, expansion rate of stalk in maturity shorten9.15and10.40%. Seedling mortality of Jijing81change less than Changbai9, low temperatureresistance level of Changbai9is from3level to1level. The result show that soil increasedphosphate fertilizer can improve cold tolerance ability in northeast japonica rice.
II．Low temperature in booting stage results in decline in seed setting rate, mature plantheight and headings of Jijing81and Changbai9. However, adding phosphate fertilizer in thebasal, mature plant heigh of two varieties increase12.4%and15.4%, headings increase0.77times and6.63times, seed setting rate of Jijing81is range from72.3~88.0%, low temperatureresistance level is from3level to1level, but seed setting rate of Changbai9is range from52.3~78.3%, low temperature resistance level of is from5level to3level, The result showthat increased phosphate fertilizer can improve cold tolerance ability in northeast japonicarice.
Ⅲ. Net photosynthetic rate of two varieties decrease79.5%and69.8%in transplantstage under low temperature, and in booting stage they decrease12.8%and19.4%.Fluorescence dynamic analysis shows that Fv/Fm decrease by adding phosphate fertilizer. ForJijing81, qP in crease and NPQ decrease, when phosphate fertilizer application is120kg/hm~2,qP reach maximum(0.70)and NPQ reach minimum(0.37). the reasults show that cold resistantvarieties of Jijing81can use higher NPQ to dissipate. For Changbai9,qP and NPQ increase,when phosphate fertilizer application is120kg/hm~2, it reach maximum. The reasults showthat Changbai9can use higher qP to reduce reflect the center of damag.but NPQ is increase,so increaseing phosphate fertilizer application can increase the radiation heat dissipation ofenergy, reduce the damage of low temperature stress, improve the northeast japonica rice coldtolerance ability.
Ⅳ. Under the low temperature, increasing phosphate fertilizer application，membranepermeability and MDA decreased，but soluble sugar content，proline, peroxidase(POD),catalase(CAT) and superoxide dismutase (SOD) increased. Compared with Jijing81, PODactivity in Changbai9is increased higher when applying more P. The activity of POD, CATand SOD are increased38.2%,39.7%and98.9%accordingly. The results indicate thatappropriate increase the amount of fertilizer P will enhance membrane permeability of leavesand the activity of POD, so that increase the cold disaster resistance of Japonica rice innortheast China.
Ⅴ. The low temperature in transplanting period causes output decrease of Japonica81and Changbai9in many aspects such as secondary branch number, ear length, panicles perhole, grains per panicle and grain weight, among which the influential order follows as grainsper panicle> panicles per hole>grain weight. However, after adding Phosphate fertilizer inthe bed soil, the above mentioned factors improve greatly for the two varieties. The lowtemperature in booting stage causes decrease of Japonica81and Changbai9in many aspectssuch as secondary branch number, ear length, panicles per hole, grains per panicle and grainweight. But different from the influential factors orders in transplanting period, the order inbooting stage follows as grains per panicle> grain weight> panicles per hole. However, afteradding phosphate fertilizer in the bed soil in booting stage, the above mentioned factors improve greatly for both varieties and phosphates exert more effects on non-chilling tolerantvarieties.
Ⅵ. Under low temperature condition, phosphate can increase the brown rice rate, milledrice rate, chalkiness, the fatty acid content and the straight-chain starch content of the two ricevarieties. It can also lower chalky grain rate and protein content. According to related analysis,the quality indicators are not closely related to phosphate levels under normal temperaturecondition. However, under low temperature condition, the high milled rice rate of Japonica81reflects its significant positive correlation with phosphate level. As to Changbai9, its brownrice rate, milled rice rate and complete milled rice rate are all in high positive correlation tophosphate level. In contrast, the protein content of both rice varieties are negatively related tophosphate amount but the fatty acid content is significantly positively related to Phosphatefertilizer amount. Other quality indicators are not significantly related to phosphate.
Ⅶ. Considering the yield and other influential factors, the optimal phosphates in bed soilranges for Japonica81and Changbai9are respectively61.69g/m~2~71.18g/m~2and60.35g/m~2~68.02g/m~2. It can reduce effect of the growing process in rice under lowtemperature stress. The optimal basic phosphate manure ranges respectively from103.65kg/hm~2~120.44kg/hm~2to91.85kg/hm~2~114.73kg/hm~2. It can improve cold toleranceability.
引文
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