温度升高和干旱对农田生态系统水碳交换动态影响的研究进展

李 倩，高阳，王洪博，王兴鹏*，杨莹攀

温度升高和干旱对农田生态系统水碳交换动态影响的研究进展

李 倩1，高阳2，王洪博1，王兴鹏1*，杨莹攀1

（1.塔里木大学 水利与建筑工程学院，新疆 阿拉尔 843300；2.中国农业科学院 农田灌溉研究所，河南 新乡 453002）

气候变化是威胁全球农业可持续发展的重要因素之一。温度升高和干旱等极端天气频发是全球气候变化的主要体现方式。温度升高显著影响土壤-作物系统的表现和功能。深入理解温度升高和干旱对农田水碳动态的影响机理，需要揭示农田生态系统功能与环境因子间的互作关系及其尺度转化效应。本文从以下几个方面综述了升温和干旱胁迫对农田生态水碳动态的影响：①全球气候变化及其影响因素分析；②温度升高、干旱胁迫以及其他气候变化因子对农田生态系统水碳动态的影响；③存在问题与未来研究方向。文献显示，世界人口增加、化石燃料燃烧碳排放量增加等致使气候变暖，在此背景下，引发的气温上升以及干旱频率增加都会对农田生态系统水碳动态有一定程度的影响。温度升高改变农田生态系统的生物量积累，同时影响作物生长和水分利用过程。此外，温度升高加快了土壤活性有机碳的转化和固存。干旱条件下会降低植物光合速率、呼吸速率以及蒸腾速率，同时，在极端干旱条件下水分利用效率也会随之降低；干旱还显著影响土壤碳转化过程与排放通量。气候变化也会伴随降水格局变化，进一步会影响土壤呼吸作用，降水增加会抑制土壤呼吸，从而减缓农田水碳循环过程。然而，目前关于增温和干旱条件下农田水碳动态的研究等方面仍存在一些不足，如对单一环境因子的研究较多，而对多重胁迫的研究较少。在今后的研究中，需加强多因素（如水分、温度等）对农田生态系统水碳动态方面的研究，以期为深入认识温度和干旱对农田生态系统的影响机理提供理论支撑。

增温；干旱；生物量；水碳循环；农田生态系统

0 引言

世界人口从1990年的52.82亿增加到2020年的75.0亿，增长率从1.74下降为1.18[1]。尽管全球人口增长率有所下降，但还是呈增加的趋势。预计到2050年，世界人口将会增加到89.1亿[1]。由于人口的增长使得人类活动产生的CO2排放量也随之增加。人类活动所产生的CO2是引起气候变化的因素之一，这对全球变暖的影响是不可逆转的[2]。

1880—2012年全球陆地平均气温升高了0.85 ℃，北半球陆地则为0.64 ℃/100 a[3-4]。中国的升温趋势为0.76 ℃/100 a，据国家气象站网获取河南省1967—2016年50年气象数据显示，当地气温呈递增趋势。温度升高不利于作物生产，其带来的负面作用会抵消大气CO2量升高对产量的促进作用[5]。相关研究指出，在低纬度地区，升温对水稻干物质的形成会有抑制作用，而在温度和CO2量升高条件下，对高纬度地区水稻生产起到双重促进作用[5]。增温对玉米产量的影响效应与水稻相类似，例如，欧洲地区玉米在增温环境下产量有随温度升高而增加的趋势[6]。

大气环境质量与全球气候的变化情况极为密切。谭雪红等[7]研究指出：在1996、1999年和2003年，人类活动对大气环境影响的综合评价指数分别为0.556、0.565和0.568，整体呈递增趋势。随着化石燃料的使用增加，CH4、N2O等有害气体的排放量也随之增加，使得大气环境的污染指数递增，这些不可避免的因素使得温室效应愈演愈烈，从而导致全球变暖[8]。

全球气候变化引起了气温升高、干旱灾害频繁、CO2量升高和降水格局变化，是威胁农业可持续发展的重要因素。Wigley等[9]根据IPCC-TAR的预测结果结合其他有关资料综合分析指出，全球平均温度从1990年到2100年将上升1.7～4.9 ℃；降水格局将发生变化，变幅为±10 %[10]。中国西部的年平均气温将升高1.7～2.3 ℃，降水增加5%～23%。据分析，CO2体积分数已经由19世纪末的280 μL/L-1增加到当今的360 μL/L-1，预计到21世纪中后期将倍增[11]。

世界人口增加、人类活动等引起气候变暖从而使CO2量、降水格局等发生变化，在此背景下，利用温度升高和干旱来分析农田系统水碳交换动态的影响，进一步为气候变暖对农田作物的影响提供支撑。

1 温度升高对农田生态系统水碳动态的影响

1.1 温度升高对农田生物量的影响

温度是改变农田生态系统生物量的因素之一。与未增温处理相比，增温提升了小麦营养、生殖生长并进期和收获期的地上生物量，但未显著影响地下部生物量；小麦在营养与生殖生长并进期增温处理的总生物量显著高于未增温处理[12]。田间水分充足的条件下，增温在有效范围内可增强土壤的微生物活性[13]；但在土壤水分不足条件下, 增温可能会抑制其活性升高, 反而会使得其活性下降[14-15]。另外，有试验得出，作物生育期会因升温而缩短，地上部生物量的积累会随温度的升高而增加，另外，增温也加快作物对地下营养物质的吸收和利用[16]。

升温促进了作物地上部分的生长发育[17]，作物生产干物质的能力也会相应地增强，植物同化CO2的能力主要受叶面积的影响[18]。因此，温度升高对植物叶片数目、面积大小以及生长速率会产生一定程度的促进作用[19]。如Wang等[20]的研究得出，增温处理会优先促进植物进行光合作用器官的生长发育。可见，温度升高可以通过促进植物的代谢和微生物的活性，来进一步增加农田生物量[12-16]。吴杨周等[21]研究表明，增温与不增温相比，冬小麦各个生育期增温显著增加了地上部生物量，同时也增加了总生物量。温度变化对植物生物量分配多少可能与同一种植物不同器官代谢速率的异同相关[22]。通常认为，升温能够加快植物的周转速率，并减少光合产物向地下的分配[23]。温度对植物生物量分配的影响大于降水影响[24]，因植物水分条件变化较快，生物量分配的能力会受到植物水分条件的限制[25]。丁乐乐等[26]和吴杨周等[27]以3 a和1 a的试验得出增温对冬小麦地上生物量的影响不一致，这主要由于增温年限不同所致。可见，在增温条件下会增加农田生态系统生物量。

1.2 温度升高对农田系统水循环的影响

温度升高是全球气候变化的一个主要体现方式，显著影响农田系统的水循环过程[28]。温度升高改变作物物候期、加剧土壤蒸发、降低冠层水汽压差（），影响作物生长和水分利用过程[29]。毛明翠等[30]指出增温会加快田间水分腾发，加大农田蒸散量。王石立等[31]认为在泰安和郑州2个地区温度升高1.5 ℃时，冬小麦生育期内潜在蒸发量分别增加24 mm和32 mm，比当前气候下多5.5%和8.5%，表明温度升高加快了农田土壤的水循环过程[32-35]。当温度升高对农田水分腾发量产生显著影响，研究其原因是温度升高改变作物物候期、土壤含水率等[29]。另外，温度控制对农田水循环的研究相对较少，极有必要深入开展关于温度升高对农田系统水循环过程的研究。

1.3 温度升高对农田系统碳循环的影响

农田碳循环系统的边界不仅限于土壤呼吸、生物量还有如肥料、杀虫剂等的使用带来的排放，另外还包括土壤CO2和CH4的排放以及秸秆还田后带来的土壤固碳，即考虑到各个环节温室气体排放及固碳的整个生命周期过程[36]。随着温度的升高，土壤CO2和CH4的排放量都呈明显增加趋势，但它们的排放量的多少还受到增温速率、幅度、土壤和微生物等其他条件的限制[37-40]。另外，Wan等[41]发现，气候变暖会改变试验因子（增温）对土壤碳排放的影响效应。增温会通过影响群落结构特征使得土壤有机碳的矿化过程发生变化以及使得土壤呼吸作用增强，进一步改变麦田中CO2的释放量[42-43]。Lu等[44]和Shi等[45]的增温试验得出，通过短期增温增加大气和土壤温度进一步促进了土壤微生物活性，同时也增加其数量，进而使土壤孔隙中的CO2排放量明显上升。Hou等[46]在华北地区农田的试验指出，在增温条件下农田土壤呼吸效应较不增温相比不显著，但延长增温时间土壤呼吸会表现出递增的趋势。

Kirschbaum等[47]研究发现，气温每上升1 ℃，土壤有机碳成分损失约10%。孔雨光等[48]研究指出，利用适度升温增强酶活性进而促进了土壤有机质的分解，加快了土壤CO2产生速率。增温使得土壤微生物代谢速率加快，酶活性增强，促进了有机碳的转化和分解，排放出更多的CO2[49-50]。李俊等[51]利用5～6 a对小麦的增温处理研究得出，在增温条件下甲烷菌种对CH4的吸收能力降低。刘艳等[52]通过模拟增温试验研究表明，在冬小麦生长季和大豆生长季增温均促进土壤CO2产生速率，但对大豆田的促进作用高于冬小麦田。增温会使叶绿素量减少、光合速率降低[53]、缩短作物生育期[54-55]、减少作物产量[54,56-59]。综上所述，温度升高对农田碳循环有促进作用。

2 干旱胁迫对农田系统水碳动态的影响

2.1 干旱胁迫对农田系统水循环的影响

气候变化条件下，干旱频率会显著增加，尤其是在干旱/半干旱地区。在农田生态系统水平上，干旱会影响群落生理特性以及结构组成，从而使得相应生态系统的光合固碳过程、蒸散耗水过程及水分利用过程发生改变[60-62]。虽然关于水分亏缺对生态系统不同层次水碳循环过程影响的研究较多，但在不同时空尺度下不同界面水碳耦合过程的系统性研究比较缺乏。

张文英[63]的试验表明，植物密度随着降水呈线性减少的趋势，农田水动态的循环及消耗速率也会减慢。干旱通过降低植物叶片气孔导度来进一步增加水分传输阻力等以减少水分损失[64]，与此同时叶片的光合速率也会随气孔导度的降低而下降[64-65]。土壤水分状况是反应农田蒸发蒸腾大小及变化的一个重要因素[66-69]。夏玉米的蒸发蒸腾研究[70-73]表明：田间土壤水分通过影响植株生长进而影响植株的棵间蒸发，在水分不足时，叶片气孔会自动关闭使得植株蒸腾速率下降，从而减少组织内部的水分散失[74-75]，此时，土壤蒸发消耗的水分也会随之减少[76]。作物会因适度缺水而保持较高的水分利用效率，而干旱较严重的条件下水分利用效率会随之降低[77-78]。干旱胁迫下叶片气孔关闭气孔导度降低，以此来减少作物植株水分损失提高蒸腾效率[79]。

综上可见，干旱会使光合固碳过程、蒸散耗水过程以及水分利用过程发生改变[60-61]，另外，减少水量会引起植物密度随着降水呈现线性减少的趋势，农田水动态的循环及消耗速率也会随之减慢[63,74-76]。

2.2 干旱胁迫对农田系统碳循环的影响

当土壤中水分不足时会影响植物体内营养物质的合成和运输，也会阻碍植物对矿质养分的吸收，而进一步抑制作物产量形成[80]。在农田生态系统中，水分不足将会阻碍作物对土壤养分的获取，影响土壤有机物的转化速率，土壤缓效养分向速效养分的转化速率也会随之减慢[81]。降水对植物生物量分配的影响与干旱的严重程度有关，在严重干旱条件下，植物响应较为强烈。吕晓敏等[82]以降水和温度为研究条件来分析其对植物碳物质和生物量的影响，发现在同一温度水平下，降水量比常规情况下减少30%，生物量会明显减小，碳物质循环也会随之减缓，而降水减少15%以内对生物量以及碳物质没有显著影响。Deng等[83]指出在水分亏缺程度不同时植物叶片生物量也有所差异，农田系统生物量的分配对短期的干旱没有明显的响应[84]。另外，降水频率和降水量的多少也会影响生物量和生产力[85]。干旱也会阻碍作物对养分的获取，影响土壤有机物的转化速率[81]，同时也会影响农田系统的生物量积累[82-84]。

一般来说，当外界环境处于正常状态且土壤水分为最大田间持水量时，土壤呼吸量会达到最大，当土壤水分过高或过低时会抑制土壤呼吸作用[86]。蔡祖聪等[87]的研究表明，农田系统中含水率对CH4氧化菌活性有一定程度的影响，当土壤含水率超过田间持水量时，其活性会随含水率的增加而受到抑制[88]。通过长期增温（5 a及以上）发现，农田土壤微生物分解能力会随着土壤含水率降低而下降，进而抑制了土壤中CO2的释放[89-91]。在水分亏缺条件下土壤有机碳量较高，无机固碳能力增强[92]。

3 其他气候因子对农田系统水碳动态的影响

3.1 CO2对农田系统水碳动态的影响

CO2是植物进行光合作用的基础，增加其浓度有利于促进植物产量和生物量的形成。随着CO2量升高，植物光合速率增加，农田系统作物耗水速率、耗水量也随之增加[93]。已有的研究表明，CO2量增加对作物生产力起促进作用[94]，例如Jablonski等[94]利用Meta分析总结了大气CO2量升高对多种植物生殖生长指标的影响，结果表明大气CO2摩尔分数升高到510～790 µmol/mol，植物在生育期内的开花数量平均增加了19%，籽实数量平均增加了16%～18%。

CO2量升高，作物叶片固碳量增加，土壤呼吸增加，但是会导致叶片气孔密度下降，气孔密度对CO2的响应存在一定的阈值效应[93]。植物光合作用随着CO2量升高而增强，在一定的量处达到峰值[95]，与此同时土壤呼吸也会随CO2量增加而增加；也有研究表明CO2量增加有利于CO2分子不断地被光合反应吸收利用，从而让更多的CO2进入叶片，来促进作物产量等[96]。植物对大气CO2量升高的响应不仅包括生物量或产量的变化，还包括糖及其他碳水化合物量变化，比如蔗糖、麦芽糖等量预计平均增加27%[97-98]。此外，当大气中CO2量增加后，与碳物质贮存和能量代谢转化相关的指标也会随之增加[99]。由相关CO2量的研究表明大气CO2摩尔分数升高到550 µmol/mol后，C3作物小麦和水稻籽粒蛋白质含量下降趋势相近，均在7.8%左右；C4作物玉米下降5%左右[100]。也有模型研究表明，大气CO2量升高对植物根系生物量多少有促进作用[101]。增加大气CO2量会提高农田作物对碳水化合物的合成、积累速度以及对矿质元素的吸收速度。有关树木的研究也指出，杨树在高CO2量环境下光合C固定能力将会增加55%左右[102]。这主要是以最大限度保证大气CO2量升高对光合C固定能力的促进作用[103]。显见，CO2量升高对农田水碳动态也有一定响应，在CO2量升高的条件下作物对碳水化合物的积累会增多，另外，碳贮存和能量代谢以及农田系统作物耗水速率也会随之增加[93,99]。

3.2 降水对农田系统水碳动态的影响

降水格局改变可通过影响作物生理生态、土壤微生物和土壤温湿度进而影响农田土壤呼吸和N2O的产生与排放。干旱区降水稀少[104]，但降水格局较非干旱区变异较大[105-106]。随着温度升高，干旱区将形成单次降水量增多、缺水间隔期延长的降水格局[107]。陈隆勋[108]和翟盘茂[109-110]等采用统计分析指出我国平均年降水量呈递减趋势。作物受到降水减少的影响，其氮代谢过程和相关酶活性的改变也会影响土壤呼吸和N2O的产生与排放[111]。Jalota等[112]发现降水可以促进棉花的作物水分生产力，但是降水过多又会破坏作物生产力；姬兴杰等[113]通过对我国北方冬小麦生育期的研究表明，除成熟期外，其他均与降水量显著负相关；张明捷等[114]研究表明，冬小麦产量与1月、6—9月降水量显著相关；陈书涛等[115]研究长三角地区温度和降水对冬小麦生育期的影响表明，试验地区降水比较充沛，所以降水不是影响该地区冬小麦生育期的关键因子。徐新创等[116]对中国近几十年来降雨强度变化趋势研究表明，在全球变暖的背景下，中国东部和南部暖湿地区强降雨事件将会增多[117]，降水还对土壤温湿度产生显著影响，进而影响土壤呼吸。综上，在气候变暖的背景下，降水格局也会发生变化，进一步会影响土壤呼吸作用，降水增加会抑制土壤呼吸，从而减缓农田水碳循环过程[111,117]。

4 存在问题与未来研究方向

由于全球气候变化影响以及试验因素的增加，进一步来研究增温和干旱对农田水碳动态的影响。增温会加剧农田系统的水循环过程，进一步加快田间水分腾发速率，加大田间蒸散量。通过升高温度以及干旱条件的试验表明，农田系统的生物量、水碳以及其他指标都会随之产生不同的影响。文献综述显示，国内外有关温度升高和干旱对农田水碳动态的影响研究已有很多，并取得了一定成果，但仍存在一些不足之处。

1）关于增温和干旱对农田土壤水碳（如何通过影响土壤微生物与土壤结构，来改变碳动态和水分运动？）以及植株水碳（如何影响气孔的碳输入和水分蒸腾间的耦合关系？）影响的研究比较欠缺。

2）虽然单因素条件下有详细的研究，但双因素（温度升高和干旱胁迫）以及多因素条件下小麦农田系统水碳动态的研究仍然缺乏。应加强多因素条件下农田土壤、作物水碳动态等方面的研究，揭示多因素条件下对农田系统水碳动态的影响。

3）关于升温和干旱对农田水碳动态影响的研究多是在单一尺度下开展的，缺乏单株-群体-区域不同尺度间的系统研究。

未来，应以分析不同地区气候变化特征和规模化节水来开展温度、水分等因子对作物各个生育期生长发育以及作物产量的影响研究，进而可以通过气候特征以及作物前期的生长状况更加准确判断农田水碳循环对环境的响应过程和作物生产变化趋势。

1）应多集中于多因素条件下对农田作物水碳的研究，如，增温和干旱条件下作物光合和蒸腾的耦合关系的响应机制等。

2）应加强未来农田水碳循环的研究，以便对区域水循环、碳循环以及地下水平衡有定量的把握，为农田系统作物生长发育进一步提供有力的支撑和依据。

3）多关注升温和干旱对农田水碳动态在单株-群体-区域不同尺度间的系统性研究，为节水高产农业发展提供理论参考。

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Changes in Water and Carbon in Farmland Ecosystems Due to the Combined Impact of Temperature Rise and Drought: A Review

LI Qian1, GAO Yang2, WANG Hongbo1, WANG Xingpeng1*, YANG Yingpan1

(1. College of Water Resource and Architecture Engineering,Tarim University,Alar 843300,China;2. Farmland Irrigation Research Institute,Chinese Academy of Agricultural Sciences,Xinxiang 453002,China)

Climate change would increase the frequency of extreme weather events threatening sustainable agricultural production as a warmed atmosphere has significant effects on functions of soil-crop systems. Although moderate drought is known to improve plant water use efficiency (), how warming combined with drought affects terrestrial ecosystems remains largely unknown. Evolutionally, different plants have developed different strategies to facilitate water acquisition, and their physiological and morphological traits respond to drought in different ways. In this paper, we reviewed the combined effects of warming and drought on water and carbon dynamics in cropped lands in three aspects: ①global climate change and its underlying drivers; ②effects of warming, drought and other climate change factors on water/carbon dynamics; ③unknown/known problems and future research directions. Global warming induced by the increased population, carbon emissions from fossil fuel and increase in drought frequency will all pose a significant impact on water and carbon in farmland. Warming will change the amount of biomass thereby affecting crop growth and water use efficiency, in addition to the accelerated soil organic carbon (SOC) loss. In the above-ground, drought will reduce photosynthetic rate and transpiration, while in the below-ground, it will reduce microbial activity and slow down SOC decomposition, thereby inhibiting soil CO2emissions. Climate change is likely to change the precipitation pattern, which in turn will alter soil respiration. Apparently, there is a lack of studies on water and carbon cycles under combined influence of drought and increasing temperature. This should be strengthened as water and carbon cycles in soil-crop systems are likely to be affected by multiple biotic and abiotic factors including soil water, temperature, carbon and nutrient concentration, in order to provide a mechanistic understanding of their combined effects on functions of farmland ecosystems as well as their feedback interaction with global warming.

global warming; drought; biomass; water and carbon cycle; farmland ecosystem

S181

A

10.13522/j.cnki.ggps.2021285

1672 - 3317（2021）12 - 0110 - 09

李倩, 高阳, 王洪博, 等. 温度升高和干旱对农田生态系统水碳交换动态影响的研究进展[J]. 灌溉排水学报, 2021, 40(12): 110-118.

LI Qian, GAO Yang, WANG Hongbo, et al. Changes in Water and Carbon in Farmland Ecosystems Due to the Combined Impact of Temperature Rise and Drought: A Review[J]. Journal of Irrigation and Drainage, 2021, 40(12): 110-118.

2021-07-07

国家自然科学基金项目（51879267）；财政部、农业农村部：现代农业产业技术体系建设专项（CARS-03）；塔里木大学校长基金项目（TDZKSS202146）；兵团财政科技计划项目（2021AA003）

李倩（1996-），女。硕士研究生，主要从事灌溉排水理论与节水灌溉技术研究。E-mail:910021332@qq.com

王兴鹏（1978-），男。教授，博士，硕士生导师，主要从事作物高效用水技术研究。E-mail: 13999068354@163.com

责任编辑：赵宇龙