昼夜不同增温对粳稻产量和品质的影响研究
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摘要
20世纪地球表面气温已经升高了0.74℃,IPCC预测21世纪末全球平均地表温度仍将上升1.8-4.0℃,模型预测到21世纪末我国平均气温可能升高2.2-4.2℃。目前,就陆地生态系统对增温响应的试验研究主要集中在自然生态系统,关于农田生态系统对昼夜不同增温的响应及其机制方面的研究很少。在研究手段上也主要是采用模型预测分析和对历史数据的分析,相关试验研究主要是在温室或开项箱中进行,很难反映作物的实际响应。水稻作为我国第一大粮食作物,约占粮食总产量的40%。研究全球变暖背景下水稻产量及品质对昼夜不同增温的响应特征及其机制,对增强人类有关全球气候变化与陆地生态系统关系的认识,降低气候变化下作物生产力预测的不确定性以及确保未来我国粮食安全具有重要的意义。为此,笔者参考国际上相关的增温系统,于2006-2008年在江苏南京应用农田开放式主动增温系统(FATI:Free Air Temperature Increase),系统研究昼夜不同增温对江淮稻区水稻生育期、地上生物量积累、产量及其构成、籽粒充实、植株光合特性、稻米品质和籽粒碳氮同化关键酶等的影响,主要结论如下：
开放式主动增温系统可以形成2 m×2 m均匀且稳定的增温范围,可使水稻全生育期冠层日均温升高幅度小于2.0℃,且不会影响田间日平均温度的变化趋势。该稻田开放式增温系统符合未来气候变暖的变化特征,能满足江淮稻区水稻系统对未来气候变暖响应与适应的试验要求。
3种增温处理对水稻的地上部生物量和产量均有降低的趋势,且全天增温、白天增温和夜间增温分别使生物量降低9.1%、10-3%和3.3%；产量降低0.9%、6.4%和6.1%。产量降低主要归因于穗粒数和粒重的下降。增温处理也降低了水稻的成穗率和干物质转运率。同时,3种增温处理明显缩短水稻从移栽到始穗的日期而不改变整个灌浆期。
全天、白天和夜间增温处理使2007年稻米的整精米率分别提高14.2%、3.6%和2.2%,而2008年显著下降,分别降低8.3%、5.7%、5.5%；两年的垩白率和垩白度均显著增加；总淀粉含量和直链淀粉含量均下降；2007年籽粒中支／直淀粉比例下降,而2008年籽粒中支／直淀粉比例显著提高,全天、白天和夜间增温处理分别提高4.6%、3.6%和4.0%。增温处理下稻米的峰值黏度、热浆黏度、崩解值和糊化温度呈上升趋势,最终黏度、消解值和回复值呈下降趋势。2007年增温处理提高了籽粒的蛋白质含量,而2008年籽粒中蛋白质含量降低。
尽管增温处理增加了水稻叶片的光合特性,比如绿叶干重所占比率和叶绿素光合色素,但对叶片的光合速率没有显著影响。同时,增温处理促进了叶片的夜间呼吸速率,白天和夜间增温分别增加了6.4%和54.5%。3种增温处理下,实际量子产量ΦPSⅡ和PSⅡ最大量子效率Fv/Fm呈下降的趋势,但初始荧光F。有增加的趋势。
昼夜不同增温处理导致强势粒的最终粒重、千粒重、最大速率时间和实灌时间均下降。此外,所有增温处理的水稻强势粒SS活性在灌浆前期均低于常规对照,灌浆后期相反。3种增温处理的水稻强势粒ADPG-PPAse活性在灌浆前期均低于常规对照,而弱势粒的酶活性在灌浆前中期均低于常规对照。所有增温处理均提高了水稻强势粒和弱势粒SSS活性,但对水稻强势粒和弱势粒SBE活性的影响不明显。直链淀粉含量可能与SS和ADPG-PPase活性有关,且主要受这两个酶灌浆前期活性大小的影响,而受SSS活性的影响较小；而支链淀粉含量可能受SBE活性的影响较大。
3种增温处理均降低了灌浆前期籽粒的GS、GOGAT活性,灌浆中后期响应不一致。蛋白质的合成与灌浆前期GS和GOGAT活性关系密切。
增温处理均提高了水稻叶片的SOD活性,全天、白天和夜间增温处理平均分别提高6.9%、5.4%和19.1%,差异均未达显著水平。夜间增温增加了叶片POD的活性,全天和白天增温增加了水稻灌浆初期和后期的POD活性,而灌浆中期则降低；全天和白天增温处理均提高了水稻叶片的CAT活性,而夜间增温使灌浆前期叶片的CAT活性上升,灌浆后期下降。3种增温处理均降低了叶片的MDA含量,全天、白天和夜间增温处理平均分别降低12.7%、15.2%和9.1%,但差异均不显著。
Global average surface air temperature has increased about 0.74℃since 100 years ago, and will still increase about 1.8-4.0℃by the end of the 21st century (IPCC), meanwhile, according to the model predicted that average air temperature might will rise about 2.2-4.2℃by the end of the 21 century in China. At present, an experimental study on the responses of terrestrial ecosystems to warming mostly focus on the natural ecosystems, however, there are few studies on the responses and mechanisms of farmland ecosystems to different diurnal warming regimes. Since most previous researches on warming effects were based on crop modeling and historical data analyses, correlated with this experiment studies have been performed under controlled conditions, such as close greenhouses and open-top chambers with air condition facilities, but they are difficult to reflect crop actural responses at crop system scale in situ. Rice is the most important cereal crop of China, produced 40% of total grain yield. A study on the response characteristics and mechanisms and of yield and quality of rice to the different diurnal warming regimes under climate warming, and it plays a key role for people to enhance the understanding of relationship between global climate change and terrestrial ecosystems, and to reduce the uncertainties of modeling predicted for crop productivity and to ensure food security under climate change in future. Therefore, based on existing in field warming facilities in the world, we applied an experimental warming facility with three warming scenarios (AW:all-day warming; DW: daytime warming; NW:nighttime warming) using Free Air Temperature Increase (FATI) technique in 2006-2008 in Nanjing city, Jiangsu province, China. Our objectives were to (1) assess the practicability of the FATI system and effects of warming on rice growth, (2) investigate the effects of the different diurnal warming regimes on growth duration, aboveground biomass accumulation, yield and its component, grain filling, photosynthesis properties of rice plant, rice quality, the activities of carbon-nitrogen assimilation related key enzymes and antioxidant enzymes and lipid peroxidation in jianghuai region. The main conclusions are as follows:
The free air temperature increased system formed 2 m×2 m of evenly and reliably warming area both at daytime and at nighttime. The daily mean temperature in the crop canopy of the different warming regimes about less than 2.0℃, and the diurnal trends of mean temperature change in the crop canopy during the whole growth duration under warming plots were all similar to those under control. Our field warming system under FATI facility can accord with the trends of future climate warming and is suitable to study the actual responses and adaptations of rice productivity to climate warming at system-level in situ in jianghuai region.
Across the three-year experimental period, field warming tended to reduce the aboveground biomass by average 9.1%,10.3% and 3.3%, and the grain yield by 0.9%, 6.4% and 6.1% in the AW, DW, and NW plots, respectively. The decreasing trend of the grain yield was mainly attributed to the warming-led negative impacts on the filled grain number and the grain weight. Compared to the un-warmed, warming tended to decrease the panicle rate and the dry matter translocation rate (DMT rate). Meanwhile, field warming led to an obvious shortness of the initial heading date, while grain filling duration almost kept unchanged.
The AW, DW and MW increased head rice rate by 14.2%,3.6% and 2.2% in 2007 and decreased significantly by 8.3%,5.7% and 5.5% in 2008, respectively. All the treatments increased significantly chalky grain rate and chalkiness of rice grain, and decreased the starch content and amylose content of rice grain during the two years. Field warming led to the reduction the ratio of amylopectin to amylose in 2007, while the increase significantly it by 4.6%,3.6% and 4.0% in the AW, DW and NW plots in 2008, respectively. There were increasing trends of peak viscosity, hot viscosity, break down and pasting temperature, and decreasing trends of final viscosity, setback and consistency in rice grain under the warmed plots. The content of grain protein was increased in 2007, while it was decreased in 2008 by warming treatments.
Although warming treatments enhanced rice photosynthetic parameters, such as green leaves biomass rate and photosynthetic pigment of flag leaf, there were no impacts on leaf photosynthetic rate. Furthermore, warming stimulated leaf respiration rate at nighttime existed in the DW and NW plots which were 6.4% and 54.5% higher than those in the control plots, respectively. There were decreasing trends of photosynthetic system II (PS II) actual photochemical efficiency (ΦPSⅡ) and maximum photochemical efficiency (Fv/Fm), while increasing trends of leaf original fluorescence (Fo).
Warming-induced a reduction of the maximum weight of a kernel,1000-grain weight, the time reaching the maximum grain rate and the time of grain filling reaching 99% of superior grain. Moreover, the activity of SS in superior grain decreased under the warmed plots during the early phase of rice grain filling, while had certain differences during the late phase of rice grain filling. The activity of ADPG-PPase in superior grain reduced under the warmed plots during the early phase of rice grain filling, while the activity of ADPG-PPase in inferior grain decreased under the warming plots during the early and middle phases of rice grain filling. Warming treatments enhanced the activities of SSS in superior and inferior grains, while had no significant effect on the activities of SBE in superior and inferior grains during the whole grain filling. The impact of SS and ADPG-PPase activity were greater on amylose content and that of SBE on amylopectin content in rice during the early phase of rice grain filling.
The activities of GS and GOGAT in grain decreased under the warmed plots during the early phase of rice grain filling, while had certain differences during the late phase of rice grain filling. The protein synthesis was closely related to the activities of GS and GOGAT during the early phase of rice grain filling.
The activity of superoxide dismatase (SOD) increased under the warmed plots, and AW, DW and NW enhanced the activity of SOD by 6.9%,5.4% and 19.1% on average during the whole grain filling. The activity of peroxidase (POD) increased in NW plot, while the activity of POD increased under the AW and DW plots during the early and late phases of rice grain filling, and it reduced during the middle phase of rice grain filling. The activity of catelase (CAT) increased under the AW and DW plots, while the activity of CAT elevated in NW plot during the early phase of rice grain filling, and it decreased during the late phase of rice grain filling. Warming treatments decreased the content of malondialdehyde (MDA), which were 12.7%,15.2% and 9.1% lower on average in the AW, DW and NW plots than the control during the whole grain filling, respectively.
开放式主动增温系统可以形成2 m×2 m均匀且稳定的增温范围,可使水稻全生育期冠层日均温升高幅度小于2.0℃,且不会影响田间日平均温度的变化趋势。该稻田开放式增温系统符合未来气候变暖的变化特征,能满足江淮稻区水稻系统对未来气候变暖响应与适应的试验要求。
3种增温处理对水稻的地上部生物量和产量均有降低的趋势,且全天增温、白天增温和夜间增温分别使生物量降低9.1%、10-3%和3.3%；产量降低0.9%、6.4%和6.1%。产量降低主要归因于穗粒数和粒重的下降。增温处理也降低了水稻的成穗率和干物质转运率。同时,3种增温处理明显缩短水稻从移栽到始穗的日期而不改变整个灌浆期。
全天、白天和夜间增温处理使2007年稻米的整精米率分别提高14.2%、3.6%和2.2%,而2008年显著下降,分别降低8.3%、5.7%、5.5%；两年的垩白率和垩白度均显著增加；总淀粉含量和直链淀粉含量均下降；2007年籽粒中支／直淀粉比例下降,而2008年籽粒中支／直淀粉比例显著提高,全天、白天和夜间增温处理分别提高4.6%、3.6%和4.0%。增温处理下稻米的峰值黏度、热浆黏度、崩解值和糊化温度呈上升趋势,最终黏度、消解值和回复值呈下降趋势。2007年增温处理提高了籽粒的蛋白质含量,而2008年籽粒中蛋白质含量降低。
尽管增温处理增加了水稻叶片的光合特性,比如绿叶干重所占比率和叶绿素光合色素,但对叶片的光合速率没有显著影响。同时,增温处理促进了叶片的夜间呼吸速率,白天和夜间增温分别增加了6.4%和54.5%。3种增温处理下,实际量子产量ΦPSⅡ和PSⅡ最大量子效率Fv/Fm呈下降的趋势,但初始荧光F。有增加的趋势。
昼夜不同增温处理导致强势粒的最终粒重、千粒重、最大速率时间和实灌时间均下降。此外,所有增温处理的水稻强势粒SS活性在灌浆前期均低于常规对照,灌浆后期相反。3种增温处理的水稻强势粒ADPG-PPAse活性在灌浆前期均低于常规对照,而弱势粒的酶活性在灌浆前中期均低于常规对照。所有增温处理均提高了水稻强势粒和弱势粒SSS活性,但对水稻强势粒和弱势粒SBE活性的影响不明显。直链淀粉含量可能与SS和ADPG-PPase活性有关,且主要受这两个酶灌浆前期活性大小的影响,而受SSS活性的影响较小；而支链淀粉含量可能受SBE活性的影响较大。
3种增温处理均降低了灌浆前期籽粒的GS、GOGAT活性,灌浆中后期响应不一致。蛋白质的合成与灌浆前期GS和GOGAT活性关系密切。
增温处理均提高了水稻叶片的SOD活性,全天、白天和夜间增温处理平均分别提高6.9%、5.4%和19.1%,差异均未达显著水平。夜间增温增加了叶片POD的活性,全天和白天增温增加了水稻灌浆初期和后期的POD活性,而灌浆中期则降低；全天和白天增温处理均提高了水稻叶片的CAT活性,而夜间增温使灌浆前期叶片的CAT活性上升,灌浆后期下降。3种增温处理均降低了叶片的MDA含量,全天、白天和夜间增温处理平均分别降低12.7%、15.2%和9.1%,但差异均不显著。
Global average surface air temperature has increased about 0.74℃since 100 years ago, and will still increase about 1.8-4.0℃by the end of the 21st century (IPCC), meanwhile, according to the model predicted that average air temperature might will rise about 2.2-4.2℃by the end of the 21 century in China. At present, an experimental study on the responses of terrestrial ecosystems to warming mostly focus on the natural ecosystems, however, there are few studies on the responses and mechanisms of farmland ecosystems to different diurnal warming regimes. Since most previous researches on warming effects were based on crop modeling and historical data analyses, correlated with this experiment studies have been performed under controlled conditions, such as close greenhouses and open-top chambers with air condition facilities, but they are difficult to reflect crop actural responses at crop system scale in situ. Rice is the most important cereal crop of China, produced 40% of total grain yield. A study on the response characteristics and mechanisms and of yield and quality of rice to the different diurnal warming regimes under climate warming, and it plays a key role for people to enhance the understanding of relationship between global climate change and terrestrial ecosystems, and to reduce the uncertainties of modeling predicted for crop productivity and to ensure food security under climate change in future. Therefore, based on existing in field warming facilities in the world, we applied an experimental warming facility with three warming scenarios (AW:all-day warming; DW: daytime warming; NW:nighttime warming) using Free Air Temperature Increase (FATI) technique in 2006-2008 in Nanjing city, Jiangsu province, China. Our objectives were to (1) assess the practicability of the FATI system and effects of warming on rice growth, (2) investigate the effects of the different diurnal warming regimes on growth duration, aboveground biomass accumulation, yield and its component, grain filling, photosynthesis properties of rice plant, rice quality, the activities of carbon-nitrogen assimilation related key enzymes and antioxidant enzymes and lipid peroxidation in jianghuai region. The main conclusions are as follows:
The free air temperature increased system formed 2 m×2 m of evenly and reliably warming area both at daytime and at nighttime. The daily mean temperature in the crop canopy of the different warming regimes about less than 2.0℃, and the diurnal trends of mean temperature change in the crop canopy during the whole growth duration under warming plots were all similar to those under control. Our field warming system under FATI facility can accord with the trends of future climate warming and is suitable to study the actual responses and adaptations of rice productivity to climate warming at system-level in situ in jianghuai region.
Across the three-year experimental period, field warming tended to reduce the aboveground biomass by average 9.1%,10.3% and 3.3%, and the grain yield by 0.9%, 6.4% and 6.1% in the AW, DW, and NW plots, respectively. The decreasing trend of the grain yield was mainly attributed to the warming-led negative impacts on the filled grain number and the grain weight. Compared to the un-warmed, warming tended to decrease the panicle rate and the dry matter translocation rate (DMT rate). Meanwhile, field warming led to an obvious shortness of the initial heading date, while grain filling duration almost kept unchanged.
The AW, DW and MW increased head rice rate by 14.2%,3.6% and 2.2% in 2007 and decreased significantly by 8.3%,5.7% and 5.5% in 2008, respectively. All the treatments increased significantly chalky grain rate and chalkiness of rice grain, and decreased the starch content and amylose content of rice grain during the two years. Field warming led to the reduction the ratio of amylopectin to amylose in 2007, while the increase significantly it by 4.6%,3.6% and 4.0% in the AW, DW and NW plots in 2008, respectively. There were increasing trends of peak viscosity, hot viscosity, break down and pasting temperature, and decreasing trends of final viscosity, setback and consistency in rice grain under the warmed plots. The content of grain protein was increased in 2007, while it was decreased in 2008 by warming treatments.
Although warming treatments enhanced rice photosynthetic parameters, such as green leaves biomass rate and photosynthetic pigment of flag leaf, there were no impacts on leaf photosynthetic rate. Furthermore, warming stimulated leaf respiration rate at nighttime existed in the DW and NW plots which were 6.4% and 54.5% higher than those in the control plots, respectively. There were decreasing trends of photosynthetic system II (PS II) actual photochemical efficiency (ΦPSⅡ) and maximum photochemical efficiency (Fv/Fm), while increasing trends of leaf original fluorescence (Fo).
Warming-induced a reduction of the maximum weight of a kernel,1000-grain weight, the time reaching the maximum grain rate and the time of grain filling reaching 99% of superior grain. Moreover, the activity of SS in superior grain decreased under the warmed plots during the early phase of rice grain filling, while had certain differences during the late phase of rice grain filling. The activity of ADPG-PPase in superior grain reduced under the warmed plots during the early phase of rice grain filling, while the activity of ADPG-PPase in inferior grain decreased under the warming plots during the early and middle phases of rice grain filling. Warming treatments enhanced the activities of SSS in superior and inferior grains, while had no significant effect on the activities of SBE in superior and inferior grains during the whole grain filling. The impact of SS and ADPG-PPase activity were greater on amylose content and that of SBE on amylopectin content in rice during the early phase of rice grain filling.
The activities of GS and GOGAT in grain decreased under the warmed plots during the early phase of rice grain filling, while had certain differences during the late phase of rice grain filling. The protein synthesis was closely related to the activities of GS and GOGAT during the early phase of rice grain filling.
The activity of superoxide dismatase (SOD) increased under the warmed plots, and AW, DW and NW enhanced the activity of SOD by 6.9%,5.4% and 19.1% on average during the whole grain filling. The activity of peroxidase (POD) increased in NW plot, while the activity of POD increased under the AW and DW plots during the early and late phases of rice grain filling, and it reduced during the middle phase of rice grain filling. The activity of catelase (CAT) increased under the AW and DW plots, while the activity of CAT elevated in NW plot during the early phase of rice grain filling, and it decreased during the late phase of rice grain filling. Warming treatments decreased the content of malondialdehyde (MDA), which were 12.7%,15.2% and 9.1% lower on average in the AW, DW and NW plots than the control during the whole grain filling, respectively.
引文
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