营林措施对森林土壤甲烷吸收的影响*

王会来 刘 娟 姜培坤 周国模 李永夫 吴家森

(浙江农林大学 亚热带森林培育国家重点实验室 浙江省森林生态系统碳循环与固碳减排重点实验室 临安 311300)



营林措施对森林土壤甲烷吸收的影响*

王会来 刘 娟 姜培坤 周国模 李永夫 吴家森

(浙江农林大学 亚热带森林培育国家重点实验室 浙江省森林生态系统碳循环与固碳减排重点实验室 临安 311300)

以期为全球气候变暖背景下的林地合理经营管理提供依据。 利用Scopus，Web of Science，SDOS，CNKI 等数据库，查询林地土壤CH4的相关文献，对不同营林措施(施肥、采伐、火烧、林下植被管理)森林土壤CH4吸收通量方面的研究进行综述。 施加N肥对于富氮森林土壤CH4吸收有抑制作用，但可以显著促进贫氮森林土壤CH4吸收； 火烧后土壤CH4吸收通量受到多种因素的影响，因此存在一定的不确定性，多数研究表明，火烧减少土壤CH4吸收通量； 皆伐改变土壤温度、含水量、有机碳的分解和利用等，从而减弱森林土壤CH4吸收能力； 择伐对森林土壤CH4吸收的影响表现为抑制、促进和无影响； 剔除林下植被提高土壤温度，加快土壤水分蒸发散失，增强CH4氧化菌的活性，促进土壤CH4吸收； 种植固氮植物使森林土壤转变为CH4的排放源。 目前经营措施对森林土壤CH4吸收影响的研究结果还存在较大差异，对营林措施影响森林土壤CH4吸收的内在机理的认识还不充分。随着研究方法和观测手段的不断发展，今后应深入研究多种因素和气候变化对林地土壤CH4吸收影响的内在机理以及甲烷氧化菌对各种干扰因素的响应机制。

人为干扰； 土壤甲烷吸收； 经营管理； 施肥； 采伐； 火烧； 林下植被管理

甲烷是引起全球气候变化的第二大温室气体，单位质量CH4的增温潜势是CO2的25倍(IPCC， 2007)，对全球气候变暖的贡献率约为20%(Duttaetal.， 2015)。截止到2011年，大气中CH4的浓度相较于工业革命前提高了150%(IPCC， 2013)，目前仍以每年0.8%的速度不断递增(IPCC， 2007)。大气中CH4通常来源于水稻田、湿地、海洋以及化石燃料的开采和燃烧，全球大气CH4年排放量为548 Tg·a-1，其中湿地、水稻田等自然排放源占30%～50%(Heimann， 2010； Kirschkeetal.， 2013)。森林土壤是大气CH4的重要吸收汇，全球土壤CH4年吸收量为 26～36 Tg·a-1，其中森林土壤CH4年吸收量占52%(Denmanetal.， 2007； Borkenetal.， 2009)。中国森林覆盖面积为2.08×108hm2，占国土总面积的22%，蕴含着巨大的CH4吸收能力。据统计，我国森林土壤CH4年吸收量为0.675 Tg·a-1，其中东部湿润、半湿润地区土壤CH4年吸收量占82%(Cai， 2012； Wangetal.， 2014)。

施肥、采伐、火烧、林下植被管理等营林措施改变了土壤含水量、土壤pH值、土壤碳含量和土壤氮含量等基本理化性质和土壤微生物的群落组成、活性等，显著影响了森林土壤CH4吸收。近年来，开展了大量营林措施对森林土壤CH4吸收影响研究，但因管理措施、森林类型、土壤状况以及气候因素的不同，营林措施对林地土壤CH4吸收影响的研究结果存在较大差异； 同一种营林措施在不同森林类型、土壤状况和气候条件下，也会产生抑制、促进和不变3种结果。这种研究结果的差异性及其作用机理的认识还不充分(Zhangetal.， 2015; Iwataetal.， 2015； Hoyos-Santillanetal.， 2016)。本文综述了营林措施(施肥、采伐、火烧和林下植被管理)影响林地土壤甲烷(CH4)吸收通量的研究进展，探讨了营林措施影响土壤CH4吸收的主要机理，并提出未来研究的重点，以期对全球气候变暖背景下林地的合理经营管理起到借鉴和启示作用。

1 施肥

施肥直接改变土壤有机碳含量和植物根系生物量，同时影响土壤微生物活性、土壤pH值等土壤理化性质，从而显著影响森林土壤CH4吸收。施肥特别是施加N肥对森林土壤CH4吸收的影响机制主要包括： 氮素累积产生的抑制作用、土壤酸化、代谢产物的毒害作用、NH4+和CH4竞争甲烷单氧酶等(程淑兰等， 2012)。施肥对林地土壤CH4吸收的影响受植被类型、施肥种类和地带气候等因素的共同影响，因此产生抑制、促进和不变3种结果，但以抑制的研究结果居多(表1)。

与此相反的是，在贫N的森林土壤中，N肥施用对土壤CH4吸收通量的影响多表现为不变或促进(表1)。贫N的寒温带森林土壤，外源氮很容易被植被和土壤微生物吸收利用(高文龙等， 2013)，而且地表长期覆盖冻土导致外源氮难以穿透到土壤CH4氧化区域(Sjögerstenetal.， 2007)，从而导致施加N肥后土壤CH4氧化速率没有显著变化(Whalenetal.， 2000)。高文龙等(2013)对寒温带针叶林和Whalen等(2000)对北方针叶林的研究表明，N肥的输入没有显著改变土壤CH4吸收能力。贫N 森林土壤由于缺乏矿质氮，土壤CH4氧化能力较弱，施加少量N肥后，Ⅰ，Ⅱ和X型甲烷氧化菌的活性增强(Aumanetal.， 2001)，从而促进林地土壤CH4的吸收。少数研究表明，短时间N肥的施加促进了土壤CH4的吸收(Veldkampetal.， 2013)。

2 火烧

火烧一方面释放大量的温室气体，造成森林生态系统碳损失，另一方面影响森林生态系统的形成与演化，改变森林生态系统的碳、氮循环过程。火烧改变土壤温度、湿度和pH值，并通过有机碳燃烧和改变黏土矿物来影响土壤结构，使得土壤CH4吸收通量发生改变(任乐等， 2014)。此外，火烧后土壤微生物以及土壤酶活性也会影响土壤CH4吸收(Zhaoetal.， 2015)。

表1 不同林分土壤CH4吸收对增氮的响应Tab.1 Responses of CH4 uptake flux to N fertilizer addition in typical stand soils

表2 不同林分土壤CH4吸收对火烧的响应Tab.2 Responses of CH4 uptake to burning in typical forest soils

3 采伐

森林采伐后，地表植被和凋落物减少，土壤裸露在地表，土壤侵蚀和淋溶作用加强，加快有机碳分解速率，减弱森林碳汇能力(闫美芳等， 2010； Zhouetal.， 2015)。采伐改变森林土壤水热条件，树木对土壤水分吸收能力的减弱使得地下水位上升，从而减弱土壤的通气透水性，进而影响森林土壤CH4吸收(Gaoetal.， 2015)。

择伐是森林采伐中最常见方式之一。择伐可以优化森林林龄结构，提高林木生产力，增加光照利用率，改善土壤水热条件，维持植物根系和微生物群落的稳定，减少林火风险，是维持森林健康的重要措施。目前，择伐对土壤CH4吸收速率影响的研究还没有统一的结论(表4)。Bradford等(2000)研究发现，温带山毛榉(Quercusrobur)人工林择伐2年后土壤CH4吸收速率增加。Yoshiyuki等(2004)发现日本扁柏(Chamaecyparisobtusa)人工林择伐8个月后土壤密度增加，土壤通气性减弱导致土壤CH4吸收通量下降。Sundqvist等(2014)发现择伐1年后土壤CH4吸收速率下降了50%。Sullivan等(2008)发现择伐1年后促进了林下植被生长，森林土壤有机碳没有显著减少，对甲烷氧化菌影响较小(Wuetal.， 2011)，土壤CH4吸收速率没有显著变化。由此可见，择伐后采样时间可能是影响土壤CH4吸收速率变化的原因之一。

4 林下植被管理

林下植被是林下生态系统的重要组成部分，林下植被管理通过改变土壤有机物的输入、小气候和土壤理化性质，对土壤CH4吸收有着重要的影响(表5)。林下种植固氮植物能增强植物根系活性，提高作物生产力，同时增加了土壤凋落物输入，促进土壤碳的积累(Wangetal.， 2014)。在土壤湿度较高的条件下，林下种植固氮植物，刺激了CH4产生菌的活性，使森林土壤转变为CH4的排放源(Qiao，etal., 2011)。Li(2010)发现种植翅荚决明(Cassiaalata)后抑制了尾叶桉(Eucalyptusurophylla)林土壤CH4吸收。

剔除尾叶桉林下灌草增大森林表层土的通透度，地表温度升高加快土壤水分蒸发散失，使土壤湿度降低刺激了甲烷氧化菌的活性，从而增加土壤CH4吸收(Wuetal., 2011; Wangetal., 2011)。刘娟等(2015)研究发现，留养杂草和剔除杂草的山核桃(Caryacathayensis)林均表现为土壤CH4的汇，剔除杂草后土壤CH4吸收通量显著增加。

除草剂改变土壤微生物数量和群落活性，进而影响土壤CH4吸收(丁洪等， 2011； 张仕颖等， 2013)。如Chen等(2009)发现，丁草胺加快了土壤甲烷吸收速率。

表3 不同林分土壤CH4吸收对皆伐的响应Tab.3 Responses of CH4 uptake to clear-cutting in typical stand soils

表4 不同林分土壤CH4吸收对择伐的响应Tab.4 Responses of CH4 uptake to thinning in typical stand soils

表5 不同林分土壤CH4吸收对林下植被管理的响应Tab.5 Responses of CH4 uptake to understory management in typical stand soils

5 展望

今后应加强以下4方面研究： 1)森林土壤CH4吸收对N输入响应机制的研究； 2)多种因素对林地土壤CH4吸收影响的长期定位试验的研究； 3)甲烷氧化菌对各种干扰因素响应模式的研究； 4)气候变化对林地土壤CH4吸收的影响及其响应机制的研究。

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(责任编辑 于静娴)

Effect of Management Practices on Methane Uptake in Forest Soils

Wang Huilai Liu Juan Jiang Peikun Zhou Guomo Li Yongfu Wu Jiasen

(State Key Laboratory of Subtropical Silviculture Zhejiang Provincial Key Laboratory of Carbon Cycling in Forest Ecosystems and Carbon Sequestration Zhejiang A & F University Lin’An 311300)

Forest soils are the main sinks of atmospheric CH4. This study is aimed to provide a basis for forest management practices under climate change. The databases of Scopus, Web of Science, SDOS and China National Knowledge Infrastructure (CNKI) were used to review current status of studies on CH4uptake from forest soils. The review highlighted the effect of forest management practices (fertilization, cutting, burning, and understory management) on soil CH4uptake, and explained its main mechanism. Fertilization tended to inhibit soil CH4uptake in N-rich forest soils, but obviously promoted soil CH4uptake in N-poor forest soils. It was generally believed that slash burning reduced soil CH4uptake. Clear-cutting changed soil temperature, water content, decomposition of organic carbon, and thus weakened the capability of methanotrophs to oxidize methane. Selective-cutting could stimulate, suppress, or show no effect on forest soil CH4uptake. Understory removal increased soil temperature, sped up evaporation of soil water and increased the activity of methanotrophs, which promoted soil CH4uptake. Nitrogen fixing plants reduced soil CH4uptake. There were significant differences of management practices on forest soil CH4uptake among the previous studies. Additionally, the inherent mechanism is still not sufficient. With the development of research theory and technology, the interactive effects of various factors and climate change on forest soil CH4uptake and its mechanism, the response model of methanotrophs to various interference factors will become the main direction in the future.Key words： human disturbance; soil CH4uptake; management practices; fertilization; cutting; burning; understory management

10.11707/j.1001-7488.20170518

2015-11-18；

2017-01-19。

浙江省自然科学基金项目(LY15C160004)； 浙江省科技创新团队项目(2012R10030-11)。

S718.5

A

1001-7488(2017)05-0156-08

* 刘娟为通讯作者。