冬小麦对昼夜不同增温的地下生物学响应特征及其机制
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摘要
自工业革命以来,全球平均气温提高了0.6～0.9℃,IPCC预测1990-2100年全球气温仍将上升1.4～5.8℃、相关预测认为2000-2050年我国平均气温仍将升高2.3～3.3℃。目前,就陆地生态系统对气候变暖响应的试验研究主要集中在自然生态系统地上部分,而地下生物学对昼夜不同增温的响应特征及其作用研究方面很少。在研究手段上也主要是模型分析预测和温室/开顶箱的试验研究,其研究结果存在较大的不确定性。冬小麦作为中国最主要的粮食作物之一,其产量占中国粮食总产的22.5%。研究全球变暖背景下冬小麦对昼夜不同增温的地下生物学响应特征及其机制对增强人类认识全球气候变化与陆地生态系统关系,降低未来气候变化预测的不确定性具有重要的意义。为此,笔者参考国际相关增温系统,于2006-2009年在江苏南京设计并运行了我国首个农田开放式主动增温系统(FATI,Free Air Temperature Increased)研究昼夜不同增温冬小麦土壤化学特性、冬小麦根系特征及活力、土壤呼吸、土壤酶学动态、土壤微生物量和土壤线虫,获得主要结论如下：
1开放式主动增温系统的增温效应显著,符合气候变暖的农田试验研究要求
开放式主动增温系统可以在2m×2m面积内增温效果显著,且不影响田间气温的变化趋势。昼夜不同增温略微降低了小麦的土壤水分,影响不显著,但显著缩短小麦生育期进程。田间结果表明本实验设计的开放式主动增温系统符合气候变暖的机制,基本能满足小麦所代表的典型农田生态系统对昼夜不同增温响应与适应的试验研究要求。
2昼夜不同增温处理降低了土壤pH值和碱解氮,土壤钾和磷有效性的响应差异明显
昼夜不同增温处理降低了土壤的pH值,影响最大为夜间增温处理,平均下降1.0%,最小为全天增温处理；增温处理也降低了各生育期土壤的碱解氮,影响规律为：夜间增温>白天增温>全天增温,小麦土壤的碱解氮分别下降3.7%、5.2%和7.8%。全天增温和白天增温处理使土壤有效磷平均分别提高5.5%和1.9%,而夜间增温却降低了2.9%。全天增温处理也使土壤的速效钾含量平均提高了10.2%。
3昼夜不同增温处理降低了小麦根冠比,提高根平均直径,根形态和活力有所变化
白天增温处理提高了根的干物质积累、根体积、根表面积、根直径、根长和根活力,对根干物质和根体积的影响达到了显著水平。夜间增温显著提高了根的干物质积累、根体积和根活力、年均分别提高10.9%、12.3%和19.3%.全天增温处理对小麦根系形态的影响存在年际差异,但差异都未达到显著水平。昼夜不同增温处理都降低了小麦孕穗-成熟的根冠比,且全天增温处理影响达到显著水平
4昼夜不同增温处理下,土壤呼吸响应明显,且夜间增温效应显著
土壤CO：日平均排放速率在越冬期前最大,越冬-拔节期最小。昼夜不同增温处理显著提高了小麦播种至拔节的土壤呼吸,提高幅度为全天增温<白天增温<夜间增温处理,拔节后增温处理对土壤呼吸的影响不显著。全生育期全天、白天增温处理表现略有下降,平均降低2.8%和2.3%,夜间增温处理使土壤呼吸了提高7.2%,差异达到显著水平。
5昼夜不同增温显著影响土壤酶活性,不同土壤酶的响应特征存在显著差异
全天增温处理使小麦各生育期土壤脲酶、蛋白酶和蔗糖酶活性年均分别提高6.3%、6.3%和2.7%；白天增温处理增加了小麦土壤脲酶和蔗糖酶活性.年均分别提高3.8%和1.2%,但降低了土壤蛋白酶活性；夜间增温处理降低了土壤的脲酶、蛋白酶和蔗糖酶活性,年均降低10.9%、16.7%和5.6%；3种增温处理都降低了小麦土壤的过氧化氢酶活性,其降低幅度为：夜间增温>白天增温>全天增温。
6昼夜不同增温降低了小麦土壤微生物量碳氮和可溶性碳氮.处理间差异显著
昼夜不同增温处理降低了小麦土壤微生物量碳和可溶性碳,影响最大为夜间增温,最小为全天增温。全天增温、白天增温和夜间增温处理分别使小麦土壤微生物量碳和可溶性碳年均降低8.4%、15.1%、29.1%和7.1%、19.2%、27.7%,小麦土壤微生物量氮和可溶性氮也表现为降低,影响最大为全天增温处理,最小为夜间增温处理。分析显示夜间增温和白天增温处理对小麦土壤微生物量碳和可溶性碳的影响达到显著水平,全天增温处理对小麦土壤微生物量氮和可溶性氮的影响也达显著水平
7昼夜不同增温处理降低了土壤线虫群体数量,改变了线虫的空间分布和营养类群
全天增温和白天增温处理使小麦0-25 cm土层线虫平均分别下降19.7%和12.6%,且全天增温处理降低达到显著水平。增温处理对0-10cm土壤线虫影响最大,全天增温和白天增温处理线虫平均分别下降37.5%和20.4%；在10-25cm土层,3种增温处理也降低了土壤线虫数量,但差异未达到显著水平。增温处理增加了0-25 cm各耕作层的食细菌线虫,略降低植食线虫,全天增温、白天增温和夜间增温处理使土壤中的食细菌线虫比例年均分别提高13.6%、18.0%和26.9%,杂食和其它食性的线虫比例有增有减,差异都未达到显著水平
Human activities and the use of fossil fuels had led the earth temperature to increase 0.6～0.9℃from the industrial revolution bingening. IPCC forecasts that the global temperatures will rise by 1.4～5.8℃from 1990 to 2100, and China's average temperature will rise by 2.3～3.3℃from 2000 to 2050 related to Chinese forecast. At present, researches of the responses of terrestrial ecosystems to experimental warming mostly focus on aboveground of natural ecosystems, while responses and adaptation of belowground biological processes to different diurnal warming are relatively scarce. Meanwhile, the main research approaches are base on modeling and the greenhouse/open-top warming methods, they cannot truely reflex crop biological processes above and below ground under the global warming. Winter wheat, as one of the most important food crops of China, produced 22.5% of total grain yield. In the context of global warming, to learn the responses of crop production to different diurnal warming scenarios and the belowground biological mechanisms plays a important role for people to enhance the understanding of relationship between global climate change and terrestrial ecosystems, and to reduce uncertainty of the future climate change prediction. Therefore, based on existing field warming facilities in the world, we designed the first Free Air Temperature Increased system for agroecosystems of China in Nanjing. Jiangsu province. In order to explore the response characteristics and mechanism of the crop belowground biological process to global warming, we conducted three diurnal warming scenarios (AW:All-daytime warming; DW:Daytime warming; NW:Nighttime warming) in the wheat field during 2007-2009. Our objectives are to (1) assess the practicability of the FATI system and effect of warming on winter wheat. (2) investigate the responses of soil physical and chemical characteristics, winter wheat root characteristics and vitality, soil and root respiration, soil enzymology dynamics, microbial biomass. soil nematodes and functional components. The main conclusions are as follows:
1 Warming effects of Free Air Temperature Increased (FATI) facility
The free air temperature increased system could significantly increase the temperature of farmland ecosystem microenvironment within the 4 m2,but did not affect the variation trend of the night field temperature. The soil water decreased slightly under different diurnal warming scenarios, but the difference did not reach significant level. Under the three different diurnal warming scenarios, the days from sowing to the beginning of earing and sowing to maturitying significantly shortened. Therefore, the free air temperature increased system adopts in the experiment accorded with climate warming mechanism, and basically satisfys the conditions which were required in the research on the responses and adaption of winter wheat to different diurnal warming scenarois.
2 Responses of soil chemical characteristics to different diurnal warming scenarios
The three diurnsl warming scenarois all reduced the soil pH. The all-day warming treatment, in which the soil pH reduced 0.6%, had the minimum effect, while night warming treatment had maximum effect, in which the difference also reached significant level in booting stage and heading stage. There was no significant effect of warming on the total soil nutrient content. Warming reduced soil alkalined-nitrogen in the whole growth and development period by 3.7%,5.2% and 7.8%. respectively, in the NW, DW and AW. All-daytime warming (AW) and Daytime warming (DW) increased soil available phosphorus by 5.5% and 1.86% respectively, however. Nighttime warming (NW) reduced by 2.9%. All-daytime warming (AW) also increased soil available potassium by average 10.2%, while Daytime warming (DW) and Nighttime warming (NW) increased soil available potassium by 3.3% and 7.5% respectively before heading stage.
3 Responses of wheat root to different diurnal warming scenarios
All-daytime warming (DW) treatments increased the root dry matter accumulation. root volume, root surface area, root diameter, root length and root activity, in which the differences of root dry matter and root volume reached significant level. Nighttime warming (NW) significantly increased root dry matter accumulation, root volume, and root activity by 10.9%.12.3% and 19.3% respectively. Wheat root morphology had inter-annual differences under All-daytime warming (AW) treatment, but did not reach significant level. All the three diurnal warming scenarois reduced the ratio of root/shoot from booting stage to maturity, in which All-daytime warming (AW) reached significant level.
4 Responses of soil respiration to different diurnal warming scenarios
Under three diurnal warming scenarios and the control, the average daily soil CO2 efflux, during the whole growth period, reached maximum before wintering period and came up to minimum during the wintering period and shooting stage. All the warming treatments significantly improved soil respiration from sowing to jointing stage, and soil respiration increased 4.9%,14.2%,23.2% for All-daytime warming (AW). Daytime warming (DW), Nighttime warming (NW) respectively. Soil respiration declined slightly for All-daytime warming (AW) and Daytime warming (DW) after jointing stage. During the whole growth period, soil respiration declined by 2.8% and 2.3% respectively for All-daytime warming (AW) and Daytime warming (DW), but did not reached significant level, however, soil respiration of Nighttime warming (NW) increased by 7.2%, and reached significant level. All the warming treatments had no effect on the daily dynamic changes of soil respiration.
5 Responses of soil enzymology to different diurnal warming scenarios
All-daytime warming (AW) treatment increased soil urease. protease and sucrase average activities by 6.3%,6.3% and 2.7% respectively. Daytime warming (DW) treatment increased two-year average activities of soil urease and sucrase by 3.8% and 1.2% respectively, but reduced the activity of soil protease. Night warming (NW) treatment yearly reduced the activities of urease, protease and sucrase by 10.9%,16.7% and 5.6% respectively. Nighttime warming (NW). Daytime warming (DW) and All-daytime warming (AW) reduced the activity of soil hydrogen peroxide by 12.3%,3.2% and 6.1% respectively. After comparing the effect of warming treatments to the activity of soil enzyme, we found that the warming effect sizes were All-daytime warming (AW)> Daytime warming (DW)> Nighttime warming (NW).
6 Responses of microbial biomass to different diurnal warming scenarios
All the warming treatments reduced the soil microbial biomass carbon and soil soluble carbon, in which Nighttime warming (NW) had maximum impact while All-Daytime warming (DW) showed the minimum effect. All-daytime warming (AW), Daytime warming (DW) and Nighttime warming (NW) treatments reduced the average soil microbial biomass carbon by 8.4%,15.1% and 29.1%, and soil soluble carbon by 7.1%, 19.2% and 27.7%, respectively. The three different diurnal warming scenarios reduced the soil microbial biomass nitrogen and soil soluble nitrogen, in which All-daytime warming (AW) had maximum impact while Nighttime warming (NW) showed minimum effect. Statistic analysis revealed that the impact of Nighttime warming (NW) and Daytime warming (DW) on the soil microbial biomass carbon and soil soluble carbon reached significant level, and only the impact of All-daytime warming (AW) to soil microbial biomass nitrogen and soil soluble nitrogen reached significant level.
7 Responses of soil nematode to different diurnal warming scenarios
All-daytime warming (AW) and Daytime warming (DW) treatment reduced the total number of nematodes which inhabited in the 0-25cm soil layer by 19.7%and 12.6% respectively. The impact of All-daytime warming (AW) on nematodes reached significant level. The warming treatments mostly affected nematodes in the 0-10cm soil layers, and All-daytime warming (AW) and Daytime warming (DW) reduced nematodes by 37.5 and 20.4% respectively. In the 10-25cm soil layer. Nighttime warming (NW). Daytime warming (DW) and All-daytime warming (AW) reduced the number of nematodes by 17.6%.14.5% and 11.4% respectively, but all the differences did not reach significant level. Warming increased the bacterivorous nematode and slightly reduced Plant-parasites All-daytime warming (AW). Daytime warming (DW) and Nighttime warming (NW) increased the numbers of nematodes averagely by 13.6%.18.0% and 26.9%. respectivlity. during the four stages, in which the booting stage and heading stage had the maximum impact. The proportions of omnivorous nematodes and other feeding habit nematodes had inconsistent variation, but none reached significant level.
1开放式主动增温系统的增温效应显著,符合气候变暖的农田试验研究要求
开放式主动增温系统可以在2m×2m面积内增温效果显著,且不影响田间气温的变化趋势。昼夜不同增温略微降低了小麦的土壤水分,影响不显著,但显著缩短小麦生育期进程。田间结果表明本实验设计的开放式主动增温系统符合气候变暖的机制,基本能满足小麦所代表的典型农田生态系统对昼夜不同增温响应与适应的试验研究要求。
2昼夜不同增温处理降低了土壤pH值和碱解氮,土壤钾和磷有效性的响应差异明显
昼夜不同增温处理降低了土壤的pH值,影响最大为夜间增温处理,平均下降1.0%,最小为全天增温处理；增温处理也降低了各生育期土壤的碱解氮,影响规律为：夜间增温>白天增温>全天增温,小麦土壤的碱解氮分别下降3.7%、5.2%和7.8%。全天增温和白天增温处理使土壤有效磷平均分别提高5.5%和1.9%,而夜间增温却降低了2.9%。全天增温处理也使土壤的速效钾含量平均提高了10.2%。
3昼夜不同增温处理降低了小麦根冠比,提高根平均直径,根形态和活力有所变化
白天增温处理提高了根的干物质积累、根体积、根表面积、根直径、根长和根活力,对根干物质和根体积的影响达到了显著水平。夜间增温显著提高了根的干物质积累、根体积和根活力、年均分别提高10.9%、12.3%和19.3%.全天增温处理对小麦根系形态的影响存在年际差异,但差异都未达到显著水平。昼夜不同增温处理都降低了小麦孕穗-成熟的根冠比,且全天增温处理影响达到显著水平
4昼夜不同增温处理下,土壤呼吸响应明显,且夜间增温效应显著
土壤CO：日平均排放速率在越冬期前最大,越冬-拔节期最小。昼夜不同增温处理显著提高了小麦播种至拔节的土壤呼吸,提高幅度为全天增温<白天增温<夜间增温处理,拔节后增温处理对土壤呼吸的影响不显著。全生育期全天、白天增温处理表现略有下降,平均降低2.8%和2.3%,夜间增温处理使土壤呼吸了提高7.2%,差异达到显著水平。
5昼夜不同增温显著影响土壤酶活性,不同土壤酶的响应特征存在显著差异
全天增温处理使小麦各生育期土壤脲酶、蛋白酶和蔗糖酶活性年均分别提高6.3%、6.3%和2.7%；白天增温处理增加了小麦土壤脲酶和蔗糖酶活性.年均分别提高3.8%和1.2%,但降低了土壤蛋白酶活性；夜间增温处理降低了土壤的脲酶、蛋白酶和蔗糖酶活性,年均降低10.9%、16.7%和5.6%；3种增温处理都降低了小麦土壤的过氧化氢酶活性,其降低幅度为：夜间增温>白天增温>全天增温。
6昼夜不同增温降低了小麦土壤微生物量碳氮和可溶性碳氮.处理间差异显著
昼夜不同增温处理降低了小麦土壤微生物量碳和可溶性碳,影响最大为夜间增温,最小为全天增温。全天增温、白天增温和夜间增温处理分别使小麦土壤微生物量碳和可溶性碳年均降低8.4%、15.1%、29.1%和7.1%、19.2%、27.7%,小麦土壤微生物量氮和可溶性氮也表现为降低,影响最大为全天增温处理,最小为夜间增温处理。分析显示夜间增温和白天增温处理对小麦土壤微生物量碳和可溶性碳的影响达到显著水平,全天增温处理对小麦土壤微生物量氮和可溶性氮的影响也达显著水平
7昼夜不同增温处理降低了土壤线虫群体数量,改变了线虫的空间分布和营养类群
全天增温和白天增温处理使小麦0-25 cm土层线虫平均分别下降19.7%和12.6%,且全天增温处理降低达到显著水平。增温处理对0-10cm土壤线虫影响最大,全天增温和白天增温处理线虫平均分别下降37.5%和20.4%；在10-25cm土层,3种增温处理也降低了土壤线虫数量,但差异未达到显著水平。增温处理增加了0-25 cm各耕作层的食细菌线虫,略降低植食线虫,全天增温、白天增温和夜间增温处理使土壤中的食细菌线虫比例年均分别提高13.6%、18.0%和26.9%,杂食和其它食性的线虫比例有增有减,差异都未达到显著水平
Human activities and the use of fossil fuels had led the earth temperature to increase 0.6～0.9℃from the industrial revolution bingening. IPCC forecasts that the global temperatures will rise by 1.4～5.8℃from 1990 to 2100, and China's average temperature will rise by 2.3～3.3℃from 2000 to 2050 related to Chinese forecast. At present, researches of the responses of terrestrial ecosystems to experimental warming mostly focus on aboveground of natural ecosystems, while responses and adaptation of belowground biological processes to different diurnal warming are relatively scarce. Meanwhile, the main research approaches are base on modeling and the greenhouse/open-top warming methods, they cannot truely reflex crop biological processes above and below ground under the global warming. Winter wheat, as one of the most important food crops of China, produced 22.5% of total grain yield. In the context of global warming, to learn the responses of crop production to different diurnal warming scenarios and the belowground biological mechanisms plays a important role for people to enhance the understanding of relationship between global climate change and terrestrial ecosystems, and to reduce uncertainty of the future climate change prediction. Therefore, based on existing field warming facilities in the world, we designed the first Free Air Temperature Increased system for agroecosystems of China in Nanjing. Jiangsu province. In order to explore the response characteristics and mechanism of the crop belowground biological process to global warming, we conducted three diurnal warming scenarios (AW:All-daytime warming; DW:Daytime warming; NW:Nighttime warming) in the wheat field during 2007-2009. Our objectives are to (1) assess the practicability of the FATI system and effect of warming on winter wheat. (2) investigate the responses of soil physical and chemical characteristics, winter wheat root characteristics and vitality, soil and root respiration, soil enzymology dynamics, microbial biomass. soil nematodes and functional components. The main conclusions are as follows:
1 Warming effects of Free Air Temperature Increased (FATI) facility
The free air temperature increased system could significantly increase the temperature of farmland ecosystem microenvironment within the 4 m2,but did not affect the variation trend of the night field temperature. The soil water decreased slightly under different diurnal warming scenarios, but the difference did not reach significant level. Under the three different diurnal warming scenarios, the days from sowing to the beginning of earing and sowing to maturitying significantly shortened. Therefore, the free air temperature increased system adopts in the experiment accorded with climate warming mechanism, and basically satisfys the conditions which were required in the research on the responses and adaption of winter wheat to different diurnal warming scenarois.
2 Responses of soil chemical characteristics to different diurnal warming scenarios
The three diurnsl warming scenarois all reduced the soil pH. The all-day warming treatment, in which the soil pH reduced 0.6%, had the minimum effect, while night warming treatment had maximum effect, in which the difference also reached significant level in booting stage and heading stage. There was no significant effect of warming on the total soil nutrient content. Warming reduced soil alkalined-nitrogen in the whole growth and development period by 3.7%,5.2% and 7.8%. respectively, in the NW, DW and AW. All-daytime warming (AW) and Daytime warming (DW) increased soil available phosphorus by 5.5% and 1.86% respectively, however. Nighttime warming (NW) reduced by 2.9%. All-daytime warming (AW) also increased soil available potassium by average 10.2%, while Daytime warming (DW) and Nighttime warming (NW) increased soil available potassium by 3.3% and 7.5% respectively before heading stage.
3 Responses of wheat root to different diurnal warming scenarios
All-daytime warming (DW) treatments increased the root dry matter accumulation. root volume, root surface area, root diameter, root length and root activity, in which the differences of root dry matter and root volume reached significant level. Nighttime warming (NW) significantly increased root dry matter accumulation, root volume, and root activity by 10.9%.12.3% and 19.3% respectively. Wheat root morphology had inter-annual differences under All-daytime warming (AW) treatment, but did not reach significant level. All the three diurnal warming scenarois reduced the ratio of root/shoot from booting stage to maturity, in which All-daytime warming (AW) reached significant level.
4 Responses of soil respiration to different diurnal warming scenarios
Under three diurnal warming scenarios and the control, the average daily soil CO2 efflux, during the whole growth period, reached maximum before wintering period and came up to minimum during the wintering period and shooting stage. All the warming treatments significantly improved soil respiration from sowing to jointing stage, and soil respiration increased 4.9%,14.2%,23.2% for All-daytime warming (AW). Daytime warming (DW), Nighttime warming (NW) respectively. Soil respiration declined slightly for All-daytime warming (AW) and Daytime warming (DW) after jointing stage. During the whole growth period, soil respiration declined by 2.8% and 2.3% respectively for All-daytime warming (AW) and Daytime warming (DW), but did not reached significant level, however, soil respiration of Nighttime warming (NW) increased by 7.2%, and reached significant level. All the warming treatments had no effect on the daily dynamic changes of soil respiration.
5 Responses of soil enzymology to different diurnal warming scenarios
All-daytime warming (AW) treatment increased soil urease. protease and sucrase average activities by 6.3%,6.3% and 2.7% respectively. Daytime warming (DW) treatment increased two-year average activities of soil urease and sucrase by 3.8% and 1.2% respectively, but reduced the activity of soil protease. Night warming (NW) treatment yearly reduced the activities of urease, protease and sucrase by 10.9%,16.7% and 5.6% respectively. Nighttime warming (NW). Daytime warming (DW) and All-daytime warming (AW) reduced the activity of soil hydrogen peroxide by 12.3%,3.2% and 6.1% respectively. After comparing the effect of warming treatments to the activity of soil enzyme, we found that the warming effect sizes were All-daytime warming (AW)> Daytime warming (DW)> Nighttime warming (NW).
6 Responses of microbial biomass to different diurnal warming scenarios
All the warming treatments reduced the soil microbial biomass carbon and soil soluble carbon, in which Nighttime warming (NW) had maximum impact while All-Daytime warming (DW) showed the minimum effect. All-daytime warming (AW), Daytime warming (DW) and Nighttime warming (NW) treatments reduced the average soil microbial biomass carbon by 8.4%,15.1% and 29.1%, and soil soluble carbon by 7.1%, 19.2% and 27.7%, respectively. The three different diurnal warming scenarios reduced the soil microbial biomass nitrogen and soil soluble nitrogen, in which All-daytime warming (AW) had maximum impact while Nighttime warming (NW) showed minimum effect. Statistic analysis revealed that the impact of Nighttime warming (NW) and Daytime warming (DW) on the soil microbial biomass carbon and soil soluble carbon reached significant level, and only the impact of All-daytime warming (AW) to soil microbial biomass nitrogen and soil soluble nitrogen reached significant level.
7 Responses of soil nematode to different diurnal warming scenarios
All-daytime warming (AW) and Daytime warming (DW) treatment reduced the total number of nematodes which inhabited in the 0-25cm soil layer by 19.7%and 12.6% respectively. The impact of All-daytime warming (AW) on nematodes reached significant level. The warming treatments mostly affected nematodes in the 0-10cm soil layers, and All-daytime warming (AW) and Daytime warming (DW) reduced nematodes by 37.5 and 20.4% respectively. In the 10-25cm soil layer. Nighttime warming (NW). Daytime warming (DW) and All-daytime warming (AW) reduced the number of nematodes by 17.6%.14.5% and 11.4% respectively, but all the differences did not reach significant level. Warming increased the bacterivorous nematode and slightly reduced Plant-parasites All-daytime warming (AW). Daytime warming (DW) and Nighttime warming (NW) increased the numbers of nematodes averagely by 13.6%.18.0% and 26.9%. respectivlity. during the four stages, in which the booting stage and heading stage had the maximum impact. The proportions of omnivorous nematodes and other feeding habit nematodes had inconsistent variation, but none reached significant level.
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