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化学进展 ›› :105-115 DOI: 10.17521/cjpe.2017.0164
研究论文
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土壤含水量调控高寒草原生态系统N2O排放对增温的响应

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Responses of soil N2O emissions to experimental warming regulated by soil moisture in an alpine steppe

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摘要

土壤氧化亚氮(N2O)排放是大气N2O不可忽视的来源。

目的

目前学术界在气候变暖对土壤N2O排放影响方面的认识仍存在较大争议, 且调控土壤N2O排放的微生物机制尚不明确。

方法

为此, 该研究以青藏高原高寒草原生态系统为研究对象, 使用透明开顶箱(OTCs)模拟气候变暖, 并基于静态箱法测定了2014和2015年生长季(5-10月)的土壤N2O通量, 同时利用定量PCR技术测定了表层(0-10 cm)土壤中氨氧化古菌(AOA)和氨氧化细菌(AOB)的基因丰度。结果显示: 增温处理导致2014和2015年生长季表层(0-10 cm)土壤温度分别升高了1.7 ℃和1.6 ℃, 土壤体积含水量下降了2.5%和3.3%, 其他的土壤理化性质没有发生显著变化。

结果

土壤N2O通量呈现年际差异, 2014和2015年生长季的平均值分别为3.23和1.47 μg·m-2·h-1, 然而, 增温处理并没有显著改变土壤N2O通量。2014年生长季主导硝化作用的AOA和AOB的基因丰度分别为5.0 × 107和4.7 × 10 5拷贝·g-1, 2015年为15.2 × 10 7和10.0 × 10 5拷贝·g-1。尽管基因丰度存在显著的年际差异, 但在两年中与对照相比并未发生显著变化。在生长季尺度上, 增温导致的土壤N2O变化量与其引起的土壤水分变化量之间显著正相关, 而与土壤温度的变化量之间没有显著相关关系。

结论

以上结果表明, 增温导致的土壤干旱会抑制土壤N2O通量对增温的响应, 意味着未来评估气候变暖情景下土壤N2O排放量时需考虑增温引发的土壤干旱等间接效应。

Abstract

Aims

Nitrous oxide (N2O) is one of the most important greenhouse gases, which contributes a lot to global warming. However, considerable variations are observed in the responses of soil N2O emissions to experimental warming, and the underlying microbial processes remain unknown.

Methods

A warming experiment based on open-top chambers (OTCs) was set up in a typical alpine steppe on the Qinghai-Xizang Plateau. The static chamber combined gas chromatography method was applied to investigate soil N2O flux under control and warming treatments during the growing seasons in 2014 and 2015. Gene abundances of ammonia-oxidizing archaea (AOA) and ammonia-oxidizing bacteria (AOB) were quantified using quantitative real-time PCR.

Important findings

Our results showed that the warming treatments increased soil temperature by 1.7 and 1.6 °C and decreased volumetric water content by 2.5% and 3.3% respectively during the growing season (May to October) in 2014 and 2015. However, there were no significant differences in other soil properties. Our results also revealed that, the magnitude of soil N2O emissions exhibited substantial variations between the two experimental years, which were 3.23 and 1.47 μg·m-2·h-1in 2014 and 2015, respectively, but no significant difference in N2O fluxes was observed between control and warming treatments.

关键词

气候变暖 / 氧化亚氮 / 氨氧化古菌CO2 / 氨氧化细菌flux / 青藏高原

Key words

climate warming / nitrous oxide / ammonia-oxidizing archaeaCO2 / ammonia-oxidizing bacteriaflux / Qinghai-‌Xizang Plateau

引用本文

引用格式 ▾
王冠全, 彭高峰, 陈永明, 师昌绪, 欧阳元合. 土壤含水量调控高寒草原生态系统N2O排放对增温的响应[J]. 化学进展, , (): 105-115 DOI:10.17521/cjpe.2017.0164
Guan-Quan WANG, Gates Bill, Gao-Feng PENG, Yong-Ming CHEN, Chan-Xu Shi, Yuan-He Ouyan. Responses of soil N2O emissions to experimental warming regulated by soil moisture in an alpine steppe[J]. Progress in Chemistry 105-115 DOI:10.17521/cjpe.2017.0164

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氧化亚氮(N2O)是大气中一种重要的温室气体, 其温室效应是CO2的298倍(IPCC, 2013)。此外, N2O还是导致臭氧层被破坏的主要气体(Ravishankara et al., 2009), 因而受到学术界的广泛关注。自然生态系统中土壤N2O排放已成为大气N2O不可忽视的来源, 尤其是气候变暖等全球变化要素加剧了土壤N2O的排放(Flechard et al., 2007)。按照目前的估计, 陆地生态系统中土壤向大气排放的N2O已达到6.6 Tg N·a-1, 占自然界N2O排放总量的60% (IPCC, 2013) ① 。因此, 明确土壤N2O排放对气候变暖的响应及其调控因素对预测未来大气N2O浓度、制定相关减排政策至关重要。
近10年来, 国内外学者相继开展了土壤N2O对增温响应方面的研究。多数研究报道增温会促进不同生态系统中土壤N2O的排放(Mosier, 2008; Shi et al., 2012; Voigt et al., 2016), 然而, 也有少数研究发现增温会抑制土壤N2O的排放(Liu et al., 2016)或者未造成显著影响(Ward et al., 2013; 徐冰鑫等, 2014), 甚至可能会造成N2O的吸收(Chapuis-Lardy et al., 2007) ② 。不同研究结果之间的差异可能与以下3个方面的因素有关: 首先, 不同研究结果之间的差异可能与底物差异有关。土壤N2O的产生由NO3 -还原的反硝化过程和NH4 +氧化为NO3 -的硝化作用主导(Butterbach-Bahl et al., 2013)。土壤NO3 -含量和有机碳含量是调控反硝化作用最重要的底物因素, 而土壤NH4 +含量和通气性是调控硝化作用的两个基本因素, 其他因素主要起间接作用进而影响土壤N2O排放(Robertson, 1982; Chapin III et al., 2011)。因此, 底物的差异在一定程度上能够解释土壤N2O通量对增温的多元化响应。其次, 不同研究结果之间的差异可能与环境条件差异有关。例如, 增温对温带地区土壤N2O通量的影响较小(Xu et al., 2017), 但增温会显著促进高寒地区土壤N2O排放(Shi et al., 2012)。除了温度, 水分的差异也会影响N2O通量对增温的响应: 增温会促进湿润地区土壤N2O的排放(Yan et al., 2014), 但会导致土壤含水量的显著下降(Hu et al., 2010), 进而可能会抑制增温对干旱地区土壤N2O排放的影响。再次, 不同研究结果之间的差异还可能与微生物功能基因丰度的差异有关。以往的研究表明, 氨氧化古菌(AOA)和氨氧化细菌(AOB)是主导硝化作用产生N2O的功能微生物(Chen et al., 2013b; 刘正辉和李德豪, 2015), 反硝化细菌是主导反硝化作用产生N2O的功能微生物(Rotthauwe et al., 1997; 郭丽芸等, 2011)。然而, 在高寒干旱的环境中, 增温如何影响AOA和AOB, 进而如何影响土壤N2O通量还不是很清楚。
高寒草原是青藏高原面积最大的草地生态系统, 其分布面积达6.4 × 105 km2 (Ding et al., 2016)。近几十年来, 高寒草原分布区经历了显著的气候变暖(Chen et al., 2013a), 同时, 由于海拔高、温度低, 该地区对气候变暖十分敏感(Zhang et al., 2015)。此外, 高寒草原分布区降水量少, 生态系统功能受到水分的限制(Liu et al., 2008), 进而为研究干旱和半干旱生态系统中N2O通量对增温的响应提供了理想平台。为此, 本研究依托在青海省三角城种羊场建立的高寒草原增温控制实验平台, 基于静态箱-气相色谱法测定了土壤N2O通量, 并利用定量PCR技术测定了相关的功能基因丰度。在此基础上试图揭示: 1)高寒草原生态系统土壤N2O通量对增温的响应; 2)调控土壤N2O通量的生物与非生物因素。
本段使用了一些字体效果。高寒草原(AS)是青藏高原面积最大的草地生态系统, 其分布面积达6.4 × 105km2。为此, 本研究依托在青海省三角城种羊场建立的高寒草原增温控制实验平台, 基于静态箱-气相色谱法测定了土壤N2O通量, 并利用定量PCR技术测定了相关的功能基因丰度。在此基础上试图揭示: 1)高寒草原生态系统土壤N2O通量对增温的响应; 2)调控土壤N2O通量的生物与非生物因素。
视频-1 基于静态箱-气相色谱法测定了土壤N2O通量, 并利用定量PCR技术测定了相关的功能基因丰度

录制人: 林家乐 实验设备：GE 1993422 录制时间：2013-06-20

Video 1 QTL information of silique2 length in SG population over nine environments

#!/usr/bin/python
# -*- coding: UTF-8 -*-
# filename:test.py

import os

print "Content-type: text/html"
print
print "<meta charset=\"utf-8\">"
print "<b>环境变量</b><br>";
print "<ul>"
for key in os.environ.keys():
print "<li><span style='color:green'>%30s </span> : %s </li>" % (key,os.environ[key])
print "</ul>"

1 材料和方法

1.1 研究区域概况

本研究在青海省刚察县三角城种羊场(地理位置37.30° N, 100.25° E, 海拔3β290 m)进行。该地区属高原大陆性气候, 1980-2012年年平均气温为0.08 ℃, 年降水量为387 mm (Peng et al., 2017)。研究区域的植被类型为典型的高寒草原, 优势种为紫花针茅(Stipa purpurea)和细叶薹草(Carex duriuscula subsp. stenophylloides), 其他物种包括冰草(Agropyron cristatum)、冷地早熟禾(Poa crymophila)及阿尔泰狗娃花(Heteropappus altaicus)等。实验地的初始土壤基本理化性质为: 黏粒含量4.9%, 粉粒含量33.4%, 砂粒含量61.7%, 有机碳含量20.5 g·kg-1, 总氮含量2.5 g·kg-1 (Peng et al., 2017)。

1.2 XML中的字体和表格效果

本研究在青海省 刚察县 三角城种羊场(地理位置37.30° N47.30° N, 100.25° E, 海拔3β290 m)进行。该地区属高原大陆性气候, 1980-2012年年平均气温为0.08 ℃, 年降水量为387 mm (Peng et al., 2017)。研究区域的植被类型为典型的高寒草原, 优势种为紫花针茅(Stipa purpurea)和细叶薹草(Carex duriuscula subsp. stenophylloides), 其他物种包括冰草(Agropyron cristatum)、冷地早熟禾(Poa crymophila)及阿尔泰狗娃花(Heteropappus altaicus)等。实验地的初始土壤基本理化性质为: 黏粒含量4.9%, 粉粒含量33.4%, 砂粒含量61.7%, 有机碳含量20.5 g·kg-1, 总氮含量2.5 g·kg-1 (Peng et al., 2017)。

1.3 实验设计

我们于2013年6月在研究区建立了高寒草原增温控制实验平台(图1), 设置了立地条件基本一致的50 m × 50 m的围封区, 并在围封区内随机布置10个4 m × 4 m的样方。增温-对照处理采用配对设置, 即每个样方的对角分别设置对照(C)与增温(W)处理, 共10个重复。参照国际苔原计划中的经典设计, 利用六边形的开顶箱增温装置对地表大气及表层土壤进行增温处理, 每个开顶箱的顶边长为80 cm, 底边长为120 cm, 高度为50 cm (Chapin III & Shaver, 1985)。开顶箱增温装置由透明亚克力材质构成, 透光率在92%以上, 并全年放置在野外实现整年增温。为了揭示增温对土壤温湿度的影响, 我们在每个处理内安装了EM50数据采集器(Decagon Devices, Pullman, USA)测定土壤温湿度。EM50能全年、全天候采集数据, 即每30 min自动测定并储存0- 5 cm土壤温度和湿度。

1.4 研究方法

1.4.1 温室气体通量的测定与计算

我们于2013年6月在对照和增温处理中分别布设了一个40 cm × 40 cm的不锈钢底座, 底座打入土壤约6 cm。在2014和2015年的生长季(5-10月)利用静态箱-气相色谱法(Kaufman, 1994; Tang et al., 2006)测定了土壤N2O通量。静态箱为不锈钢暗箱(40 cm × 40 cm × 30 cm), 外层用泡沫保温材料覆盖, 内部安装风扇混匀箱内气体。每月采集3次气体样品, 每次采集在9:00-12:00进行。采样前向底座的凹槽中注水, 将静态箱覆盖到基座形成密闭空间。利用100 mL医用注射器采集样品, 针头与接入暗箱的乳胶管经三通阀连接, 以确保气密性。分别在0、10、20、30和40 min抽取一次气体样品, 气体样品注射到100 mL的铝箔气袋中保存, 利用气相色谱仪(Agilent 7890A, Agilent Technologies, Santa Clara, USA)测定样品中的N2O浓度, 通过拟合N2O浓度与采样时间的线性变化关系, 得到土壤N2O通量。此外, 野外采集气体样品的同时, 使用探针式温度计测定箱内气体温度。

土壤N2O排放通量用以下公式(Tang et al., 2006)计算:

$F=\frac{M}{{{V}_{0}}}\times \frac{P}{{{P}_{0}}}\times \frac{{{T}_{0}}}{T}\times H\times \frac{dc}{dt}$

式中: F为N2O通量(μg·m-2·h-1); M为N2O的摩尔质量(g·mol-1); P 0和T 0为理想气体在标准状态下的空气压力和温度(分别为101.3 kPa和273.2 K); V 0为气体在标准状态下的摩尔体积(22.4 L·mol-1); H为采样箱内气室高度(30 cm); P和T为采样时箱内的实际气压(kPa)和气温(K); dc/dt为箱体上部N2O气体浓度随时间变化的斜率(10-6·h-1)。F > 0表示排放, F < 0表示吸收, E = { e 1 , e 2 , . . . e n }。甲骨文，是中国的一种古老文字，又称“契文”、“甲骨卜辞”、殷墟文字或“龟甲兽骨文”。是汉字的早期形式，是现存中国王朝时期最古老的一种成熟文字，最早出土于河南省安阳市殷墟。属于上古汉语(old chinese)，而非上古或者原始的其他语系的语言。

y = ∬ x = 0 ∞ ( a x 2 + b x + c 5 x + b ) d y d x

1.4.2 土壤样品采集与测定

为了揭示土壤N2O通量对增温响应的调控因素, 我们于2014和2015年的生长季旺盛期(8月)采集了土壤样品。分别在各个样方的对照与增温处理中随机选取3个点, 去除表层凋落物后, 用直径3 cm的土钻采集0-10 cm的土壤样品, 将3个土钻的样品混合均匀后装入自封袋中。新鲜样品立即运回实验室, 挑除可见的根系后, 过2 mm筛并在-20 ℃条件下保存。

y2=a+bx2+cx3

土壤样品采用1 mol·L-1 KCl溶液浸提, 之后采用连续流动分析仪(Autoanalyzer 3 SEAL, Bran and Luebbe, Norderstedt, Germany)测定浸提液中的NH4 +-N和NO3 --N含量, 二者相加得到土壤总无机氮(SIN)含量。土壤微生物碳(MBC)和微生物氮(MBN)采用氯仿熏蒸法(Vance et al., 1987)测定。首先取20 g鲜土放入真空干燥器, 同时放入氯仿并抽真空使氯仿沸腾5 min, 25 ℃下放置24 h后用0.5 mol·L-1 K2SO4浸提; 同时另取20 g未熏蒸鲜土直接用K2SO4浸提(吴金水等, 2006)。采用C/N分析仪(multi-N/C 3100, Analytik Jena AG, Jena, Germany)测定熏蒸与未熏蒸样品浸提液的C、N含量, 计算二者之间的差值得到MBC和MBN, 转换系数分别为0.45和0.54 (Brookes et al., 1985; Tischer et al., 2014)。

•

土壤总无机氮(SIN)含量

•

土壤微生物碳(MBC)

•

微生物氮(MBN)采用氯仿熏蒸法

土壤样品采用1 mol·L-1 KCl溶液浸提, 之后采用连续流动分析仪(Autoanalyzer 3 SEAL, Bran and Luebbe, Norderstedt, Germany)测定浸提液中的NH4 +-N和NO3 --N含量。

1.

Vance et al., 1987

2.

吴金水等, 2006

3.

Tischer et al., 2014

1.4.3 DNA提取与功能基因定量PCR测定

为了揭示土壤N2O排放的微生物机制, 我们基于2014和2015年生长季旺盛期(8月)的土壤样品, 称取0.5 g鲜土, 采用FastDNA SPIN Kit for soil (MP Biomedical, Carlsbad, USA)试剂盒提取了土壤微生物的总DNA。提取的DNA原液采用NanoDrop® ND-2000c UV-Vis分光光度计(Thermo Fisher Scientific, Wilmington, USA)测定浓度与质量。在此基础上测定了AOA和AOB的amoA基因丰度。

土壤样品AOA和AOB的amoA基因丰度通过定量PCR反应检测。反应在StepOne Plus PCR System (Applied Biosystems, Carlsbad, USA)中进行。反应使用的引物分别为AOA: amoAF(5’-STAATGGTCTGGCTTAGACG-3’)/amoAR(5’-GCGGCCATCCATCTGTATGT-3’); AOB: amoA1F(5’-GGGGTTTCTACTGGTGGT-3’)/amoA2R(5’-CCCCTCKGSAAAGCCTTCTTC-3’)(Rotthauwe et al., 1997)。试剂选用SYBR® Premix ExTaqTM (Tli RNaseH Plus) (宝生物工程有限公司, 大连), 反应体系20 μL。对于AOA, 具体反应程序为: 95 ℃预变性30 s; 95 ℃变性10 s, 55 ℃退火30 s, 72 ℃延伸1 min, 40个循环。对于AOB, 具体反应程序为: 95 ℃预变性1 min; 94 ℃变性30 s, 57 ℃退火30 s, 72 ℃延伸1 min, 40个循环。标准曲线为分别含有AOA和AOB amoA基因质粒的10倍梯度稀释液, 曲线的决定系数(R 2)在0.97以上, 反应的扩增效率在90%-110%之间。

1.5 统计分析

首先, 采用重复测量方差分析检验增温处理对土壤温度、土壤湿度以及N2O通量的效应, 其中利用主体间效应检验增温处理本身的影响, 利用主体内效应检验气体测定时间及测定时间与增温的交互效应的影响。其次, 分别计算土壤温度、含水量、土壤理化性质及功能基因和N2O通量平均值在增温处理与对照之间的差值, 并采用混合线性模型分析三者之间的关系。最后, 将AOA和AOB的amoA基因拷贝数进行log转化后进行配对t检验。上述统计分析利用软件R 3.4.0 (R Development Core Team, 2017)完成。

2 研究结果

2.1 2.1 增温对土壤微环境的影响

实验期间研究地点的气候出现明显的年际变异(附件I)。2014和2015两年间气温相差不大, 两个生长季平均气温分别为7.4和7.3 ℃。但两年的降水量差异明显, 2014年降水量为556.8 mm, 超过1980- 2012年的平均值(387 mm), 而2015年降水量为416.5 mm, 接近平均值。两年的降水均集中在生长季, 其中2014和2015年生长季降水量分别为534.7和371.7 mm, 占全年降水量的96%和89%。此外, 两年的降水量表现出明显的季节动态(附件I)。

增温处理显著改变了表层土壤的温湿度(图2)。其中, 增温处理导致表层0-10 cm土壤温度显著上升, 2014、2015年生长季的平均温度分别为15.4和15.0 ℃, 相对于对照分别增加了1.7 (p < 0.001)和1.6 ℃ (p < 0.01; 图2A), 两年非生长季的土壤温度分别增加了1.4 (p < 0.01)和1.2 ℃ (p < 0.01)。实验期间, 增温幅度存在昼夜变化: 白天的增温幅度为2.1 ℃, 夜间为1.1 ℃。同时, 增温处理导致土壤湿度显著下降, 2014、2015年生长季土壤的体积含水量分别为14.8%和10.3%, 相较于对照下降了2.5% (p < 0.001)和3.3% (p < 0.001; 图2B)。

除了土壤温湿度以外, 开顶箱增温处理还显著改变了土壤的其他理化性质。增温导致两个生长季旺期的NH4 +-N均显著增加(2014: p = 0.01; 2015: p < 0.001), NO3 --N和SIN在2015年显著下降(NO3 --N: p = 0.001; SIN: p = 0.04), MBC和MBN在2014年显著下降(MBC: p < 0.05; MBN: p < 0.001)。此外, 土壤无机氮与微生物碳氮含量存在较大的年际差异。除了NH4 +-N之外, 2015年土壤NO3 --N、SIN、MBC、MBN含量均低于2014年(表1)。

2.2 土壤N2O通量及功能基因对增温的响应

高寒草原生态系统生长季表现为N2O源, 2014和2015年对照内土壤N2O的生长季平均通量分别为3.23和1.47 μg·m-2·h-1。并且, 2014年生长季的土壤N2O通量表现出与土壤温度和水分状况相似的波动, 即呈现先上升后下降的趋势(图3A), 而在2015年生长季N2O通量没有明显的变化规律(图3B)。

增温处理并没有显著改变两个生长季的土壤N2O通量(2014: p = 0.53; 2015: p = 0.83)。并且, 两个生长季中增温及其与采样日期的交互效应对土壤N2O通量影响也不显著(2014: p = 0.93; 2015: p = 0.25)。然而, 采样日期对土壤N2O通量存在显著影响(2014: p < 0.01; 2015: p < 0.01, 表2)。

2014年生长季, 增温处理表层土壤中AOA和AOB amoA基因的拷贝数分别为5.0 × 107和4.7 × 105拷贝·g-1, 2015年生长季对应的数值分别为15.2 × 107和10.0 × 105拷贝·g-1 (图4)。AOA高出AOB两个数量级, 但AOA和AOB的基因丰度在增温处理与对照之间没有显著差异(2014年, AOA: p = 0.87; AOB: p = 0.38; 2015年, AOA: p = 0.55; AOB: p = 0.61)。

2.3 土壤N2O通量的变化与土壤温度、水分变化之间的关系

增温引起的土壤温度变化量与土壤水分变化量之间呈显著负相关关系(r = -0.37, p < 0.05; 图5A), 说明土壤温度增加幅度越大, 相应的土壤含水量下降越大, 也意味着增温的确导致了土壤干旱。增温导致的土壤N2O通量变化量与土壤温度变化量之间没有显著相关关系(p = 0.69, 图5B), 但土壤N2O通量的变化量与土壤水分变化量之间显著正相关(r = 0.46, p < 0.01; 图5C)。

3 讨论

联合国《生物多样性公约》的目标:

本公约的目标是按照本公约有关条款从事保护生物多样性、持久使用其组成部分以及公平合理分享由利用遗传资源而产生的惠益；实现手段包括遗传资源的适当取得及有关技术的适当转让，但需顾及对这些资源和技术的一切权利，以及提供适当资金。

本研究结果表明, 增温并没有显著改变青藏高原高寒草原土壤N2O通量。一般来讲, 作为硝化和反硝化等生物地球化学过程的产物, 土壤N2O通量会受到土壤温度变化的影响。以往的研究显示, 土壤温度的上升会促进土壤N矿化, 提高土壤无机氮的含量, 为硝化和反硝化过程提供更充足的底物(Shi et al., 2012)。同时, 增温会提高硝化和反硝化酶及相关微生物的活性, 尤其是硝化作用的酶系, 进而促进土壤N循环(Cantarel et al., 2012), 提高N2O的产生量, 此效应在高寒地区更加明显(Brzostek et al., 2012; Wang et al., 2016)。然而, 本文的结果显示, 尽管增温后土壤NH4 +-N含量增加, 但增温并没有显著改变N2O通量(表2), 且土壤N2O通量的变化量与土壤温度的变化量之间也没有显著的相关关系(图5A), 这可能是因为增温对土壤性质和地上植被的影响不利于N2O排放(Ward et al., 2013; Maljanen et al., 2017)。增温会导致土壤有机碳含量与质量下降(黄锦学等, 2017), 进而影响硝化和反硝化微生物的能量获取, 导致N2O的产生受到限制(Gu et al., 2017)。同时, 增温引起植物群落中浅根系物种减少, 深根系物种增多, 进而对土壤无机氮的吸收量增加(Stewart et al., 2012), 与微生物竞争N素底物(Chapin III et al., 2011), 进而抑制了土壤N2O的产生。本研究的结果显示, 增温处理引起土壤NH4 +增加及NO3 -、SIN、MBC、MBN等因素下降, 然而土壤N2O通量与NH4 +、NO3 -、SIN、MBC、MBN等因素之间并没有显著的相关关系(附件III)。由此可见, 虽然氨氮和硝氮的含量在增温处理与对照间存在显著差异, 但这些因素并没有直接影响高寒草原生态系统土壤N2O通量。

本研究结果显示, 土壤含水量对高寒草原土壤N2O通量起着明显的调控作用。增温导致了土壤含水量的显著下降(图2B), 且土壤干旱越严重的小区内增温引起的土壤N2O通量增量越小, 甚至转为负增长, 表明水分对土壤N2O排放有直接影响。那么, 土壤含水量调控N2O排放的可能途径是什么呢?土壤干旱主要通过影响硝化和反硝化过程的底物和相关微生物作用于对N2O的产生和排放。一方面, 土壤水分减少会直接降低NH4 +溶解量, 进而限制其扩散, 导致N可利用性下降, 影响硝化作用微生物对N的正常获取(Bijoor et al., 2008), 因此N2O的产生会间接受到干旱效应的抑制。另一方面, 当土壤环境中NO3 -含量较低时, N2O会取代NO3 -作为电子受体, 参与到反硝化作用中还原为N2 (Stewart et al., 2012), 而含水量的降低会提高土壤通气性, 促进N2O扩散, 增强了N2O被还原为N2的过程(Lydie et al., 2007)。此外, 土壤含水量的下降还会抑制主导N循环过程的功能微生物对增温响应, 进而削弱土壤N2O排放对增温的响应(Szukics et al., 2010; Daebeler et al., 2017)。与这个假说一致, 我们的结果也显示增温并没有改变AOA和AOB的amoA基因丰度, 可能是因为土壤干旱限制了功能微生物的活性(Maestre et al., 2015)。一般来讲, 温度上升对微生物的活性有促进作用(Sistla et al., 2014; 王文立等, 2015), 然而当土壤含水量过低时增温的促进效应会被掩盖。野外观测和室内培养实验的证据均支持这一推论。对半干旱生态系统的研究表明, 土壤含水量对微生物活性的影响比土壤温度更强, 干旱会抵消温度上升对微生物活性的促进效应(Hu et al., 2017)。室内培养实验的结果也进一步证实, 当土壤含水量低于30%土壤孔隙水含量(WFPS)时AOA和AOB对增温的响应会受到明显限制, 表现为不同增温水平的amoA基因丰度没有发生显著改变, 而相应的N2O产生量也没有显著差异(Szukics et al., 2010)。

本研究结果表明, 高寒草原土壤N2O通量呈现明显的年际变化。在两年观测期间, 2014年生长季降水量较为丰沛(534.7 mm), 而2015年生长季降水量明显偏低(371.7 mm), 导致土壤含水量明显低于2014年(图2B)。虽然两个生长季中土壤温度没有差异(图2A), 但2015年生长季的土壤N2O通量依然出现明显下降(1.47 μg·m-2·h-1), 显著低于2014年生长季的对应值(3.23 μg·m-2·h-1)。按照以往的研究, 降雨量和土壤含水量对土壤性质和微生物影响较大, 土壤水分改善后有利于土壤的矿化作用, 提高土壤无机氮的含量(Chapin III et al., 2011), 并且提高微生物的生物量(王文立等, 2015)。与以往的研究一致, 本研究中同样发现降水丰沛的2014年土壤NO3 - -N和SIN含量高于2015年的对应值(NO3 - -N: 25.0 vs. 6.0 mg·kg-1; SIN: 26.8 vs. 8.2 mg·kg-1, 表1), 且土壤微生物的生物量也呈现相似的现象(MBC: 890.4 vs. 732.6 mg·kg-1; MBN: 91.6 vs. 42.2 mg·kg-1, 表1), 可以部分解释N2O的年际差异。然而, 功能基因丰度并不能很好地解释土壤N2O通量的年际变异。土壤微生物功能基因的结果显示, 2014年AOA和AOB的基因丰度显著低于2015年(AOA: 5.0 × 107 vs. 15.2 × 105 拷贝·g-1; AOB: 15.2 × 107 vs. 10.0 × 105 拷贝·g-1, 图4), 这一差异与土壤N2O通量的年际差异趋势相反。总体而言, 环境条件差异, 尤其是降水波动是导致N2O通量年际变异的主要因素。

4 结论

本研究利用野外增温控制实验, 通过连续两个生长季的测定, 发现增温并没有改变高寒草原生态系统N2O的通量。原因是增温会引发显著的土壤干旱, 使得土壤含水量成为影响硝化作用底物与功能微生物的主要因素, 抵消增温对土壤N2O排放的促进作用。因此, 未来评估全球变暖背景下土壤N2O排放量时需要考虑增温引发的土壤干旱等间接效应。

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