營林措施對森林土壤甲烷吸收的影響*

王會來 劉 娟 姜培坤 周國模 李永夫 吳家森

(浙江農(nóng)林大學 亞熱帶森林培育國家重點實驗室 浙江省森林生態(tài)系統(tǒng)碳循環(huán)與固碳減排重點實驗室 臨安 311300)

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營林措施對森林土壤甲烷吸收的影響*

王會來 劉 娟 姜培坤 周國模 李永夫 吳家森

(浙江農(nóng)林大學 亞熱帶森林培育國家重點實驗室 浙江省森林生態(tài)系統(tǒng)碳循環(huán)與固碳減排重點實驗室 臨安 311300)

以期為全球氣候變暖背景下的林地合理經(jīng)營管理提供依據(jù)。 利用Scopus，Web of Science，SDOS，CNKI 等數(shù)據(jù)庫，查詢林地土壤CH4的相關文獻，對不同營林措施(施肥、采伐、火燒、林下植被管理)森林土壤CH4吸收通量方面的研究進行綜述。 施加N肥對于富氮森林土壤CH4吸收有抑制作用，但可以顯著促進貧氮森林土壤CH4吸收； 火燒后土壤CH4吸收通量受到多種因素的影響，因此存在一定的不確定性，多數(shù)研究表明，火燒減少土壤CH4吸收通量； 皆伐改變土壤溫度、含水量、有機碳的分解和利用等，從而減弱森林土壤CH4吸收能力； 擇伐對森林土壤CH4吸收的影響表現(xiàn)為抑制、促進和無影響； 剔除林下植被提高土壤溫度，加快土壤水分蒸發(fā)散失，增強CH4氧化菌的活性，促進土壤CH4吸收； 種植固氮植物使森林土壤轉(zhuǎn)變?yōu)镃H4的排放源。 目前經(jīng)營措施對森林土壤CH4吸收影響的研究結果還存在較大差異，對營林措施影響森林土壤CH4吸收的內(nèi)在機理的認識還不充分。隨著研究方法和觀測手段的不斷發(fā)展，今后應深入研究多種因素和氣候變化對林地土壤CH4吸收影響的內(nèi)在機理以及甲烷氧化菌對各種干擾因素的響應機制。

人為干擾； 土壤甲烷吸收； 經(jīng)營管理； 施肥； 采伐； 火燒； 林下植被管理

甲烷是引起全球氣候變化的第二大溫室氣體，單位質(zhì)量CH4的增溫潛勢是CO2的25倍(IPCC， 2007)，對全球氣候變暖的貢獻率約為20%(Duttaetal.， 2015)。截止到2011年，大氣中CH4的濃度相較于工業(yè)革命前提高了150%(IPCC， 2013)，目前仍以每年0.8%的速度不斷遞增(IPCC， 2007)。大氣中CH4通常來源于水稻田、濕地、海洋以及化石燃料的開采和燃燒，全球大氣CH4年排放量為548 Tg·a-1，其中濕地、水稻田等自然排放源占30%～50%(Heimann， 2010； Kirschkeetal.， 2013)。森林土壤是大氣CH4的重要吸收匯，全球土壤CH4年吸收量為 26～36 Tg·a-1，其中森林土壤CH4年吸收量占52%(Denmanetal.， 2007； Borkenetal.， 2009)。中國森林覆蓋面積為2.08×108hm2，占國土總面積的22%，蘊含著巨大的CH4吸收能力。據(jù)統(tǒng)計，我國森林土壤CH4年吸收量為0.675 Tg·a-1，其中東部濕潤、半濕潤地區(qū)土壤CH4年吸收量占82%(Cai， 2012； Wangetal.， 2014)。

施肥、采伐、火燒、林下植被管理等營林措施改變了土壤含水量、土壤pH值、土壤碳含量和土壤氮含量等基本理化性質(zhì)和土壤微生物的群落組成、活性等，顯著影響了森林土壤CH4吸收。近年來，開展了大量營林措施對森林土壤CH4吸收影響研究，但因管理措施、森林類型、土壤狀況以及氣候因素的不同，營林措施對林地土壤CH4吸收影響的研究結果存在較大差異； 同一種營林措施在不同森林類型、土壤狀況和氣候條件下，也會產(chǎn)生抑制、促進和不變3種結果。這種研究結果的差異性及其作用機理的認識還不充分(Zhangetal.， 2015; Iwataetal.， 2015； Hoyos-Santillanetal.， 2016)。本文綜述了營林措施(施肥、采伐、火燒和林下植被管理)影響林地土壤甲烷(CH4)吸收通量的研究進展，探討了營林措施影響土壤CH4吸收的主要機理，并提出未來研究的重點，以期對全球氣候變暖背景下林地的合理經(jīng)營管理起到借鑒和啟示作用。

1 施肥

施肥直接改變土壤有機碳含量和植物根系生物量，同時影響土壤微生物活性、土壤pH值等土壤理化性質(zhì)，從而顯著影響森林土壤CH4吸收。施肥特別是施加N肥對森林土壤CH4吸收的影響機制主要包括： 氮素累積產(chǎn)生的抑制作用、土壤酸化、代謝產(chǎn)物的毒害作用、NH4+和CH4競爭甲烷單氧酶等(程淑蘭等， 2012)。施肥對林地土壤CH4吸收的影響受植被類型、施肥種類和地帶氣候等因素的共同影響，因此產(chǎn)生抑制、促進和不變3種結果，但以抑制的研究結果居多(表1)。

與此相反的是，在貧N的森林土壤中，N肥施用對土壤CH4吸收通量的影響多表現(xiàn)為不變或促進(表1)。貧N的寒溫帶森林土壤，外源氮很容易被植被和土壤微生物吸收利用(高文龍等， 2013)，而且地表長期覆蓋凍土導致外源氮難以穿透到土壤CH4氧化區(qū)域(Sj?gerstenetal.， 2007)，從而導致施加N肥后土壤CH4氧化速率沒有顯著變化(Whalenetal.， 2000)。高文龍等(2013)對寒溫帶針葉林和Whalen等(2000)對北方針葉林的研究表明，N肥的輸入沒有顯著改變土壤CH4吸收能力。貧N 森林土壤由于缺乏礦質(zhì)氮，土壤CH4氧化能力較弱，施加少量N肥后，Ⅰ，Ⅱ和X型甲烷氧化菌的活性增強(Aumanetal.， 2001)，從而促進林地土壤CH4的吸收。少數(shù)研究表明，短時間N肥的施加促進了土壤CH4的吸收(Veldkampetal.， 2013)。

2 火燒

火燒一方面釋放大量的溫室氣體，造成森林生態(tài)系統(tǒng)碳損失，另一方面影響森林生態(tài)系統(tǒng)的形成與演化，改變森林生態(tài)系統(tǒng)的碳、氮循環(huán)過程?；馃淖兺寥罍囟?、濕度和pH值，并通過有機碳燃燒和改變黏土礦物來影響土壤結構，使得土壤CH4吸收通量發(fā)生改變(任樂等， 2014)。此外，火燒后土壤微生物以及土壤酶活性也會影響土壤CH4吸收(Zhaoetal.， 2015)。

表1 不同林分土壤CH4吸收對增氮的響應Tab.1 Responses of CH4 uptake flux to N fertilizer addition in typical stand soils

表2 不同林分土壤CH4吸收對火燒的響應Tab.2 Responses of CH4 uptake to burning in typical forest soils

3 采伐

森林采伐后，地表植被和凋落物減少，土壤裸露在地表，土壤侵蝕和淋溶作用加強，加快有機碳分解速率，減弱森林碳匯能力(閆美芳等， 2010； Zhouetal.， 2015)。采伐改變森林土壤水熱條件，樹木對土壤水分吸收能力的減弱使得地下水位上升，從而減弱土壤的通氣透水性，進而影響森林土壤CH4吸收(Gaoetal.， 2015)。

擇伐是森林采伐中最常見方式之一。擇伐可以優(yōu)化森林林齡結構，提高林木生產(chǎn)力，增加光照利用率，改善土壤水熱條件，維持植物根系和微生物群落的穩(wěn)定，減少林火風險，是維持森林健康的重要措施。目前，擇伐對土壤CH4吸收速率影響的研究還沒有統(tǒng)一的結論(表4)。Bradford等(2000)研究發(fā)現(xiàn)，溫帶山毛櫸(Quercusrobur)人工林擇伐2年后土壤CH4吸收速率增加。Yoshiyuki等(2004)發(fā)現(xiàn)日本扁柏(Chamaecyparisobtusa)人工林擇伐8個月后土壤密度增加，土壤通氣性減弱導致土壤CH4吸收通量下降。Sundqvist等(2014)發(fā)現(xiàn)擇伐1年后土壤CH4吸收速率下降了50%。Sullivan等(2008)發(fā)現(xiàn)擇伐1年后促進了林下植被生長，森林土壤有機碳沒有顯著減少，對甲烷氧化菌影響較小(Wuetal.， 2011)，土壤CH4吸收速率沒有顯著變化。由此可見，擇伐后采樣時間可能是影響土壤CH4吸收速率變化的原因之一。

4 林下植被管理

林下植被是林下生態(tài)系統(tǒng)的重要組成部分，林下植被管理通過改變土壤有機物的輸入、小氣候和土壤理化性質(zhì)，對土壤CH4吸收有著重要的影響(表5)。林下種植固氮植物能增強植物根系活性，提高作物生產(chǎn)力，同時增加了土壤凋落物輸入，促進土壤碳的積累(Wangetal.， 2014)。在土壤濕度較高的條件下，林下種植固氮植物，刺激了CH4產(chǎn)生菌的活性，使森林土壤轉(zhuǎn)變?yōu)镃H4的排放源(Qiao，etal., 2011)。Li(2010)發(fā)現(xiàn)種植翅莢決明(Cassiaalata)后抑制了尾葉桉(Eucalyptusurophylla)林土壤CH4吸收。

剔除尾葉桉林下灌草增大森林表層土的通透度，地表溫度升高加快土壤水分蒸發(fā)散失，使土壤濕度降低刺激了甲烷氧化菌的活性，從而增加土壤CH4吸收(Wuetal., 2011; Wangetal., 2011)。劉娟等(2015)研究發(fā)現(xiàn)，留養(yǎng)雜草和剔除雜草的山核桃(Caryacathayensis)林均表現(xiàn)為土壤CH4的匯，剔除雜草后土壤CH4吸收通量顯著增加。

除草劑改變土壤微生物數(shù)量和群落活性，進而影響土壤CH4吸收(丁洪等， 2011； 張仕穎等， 2013)。如Chen等(2009)發(fā)現(xiàn)，丁草胺加快了土壤甲烷吸收速率。

表3 不同林分土壤CH4吸收對皆伐的響應Tab.3 Responses of CH4 uptake to clear-cutting in typical stand soils

表4 不同林分土壤CH4吸收對擇伐的響應Tab.4 Responses of CH4 uptake to thinning in typical stand soils

表5 不同林分土壤CH4吸收對林下植被管理的響應Tab.5 Responses of CH4 uptake to understory management in typical stand soils

5 展望

今后應加強以下4方面研究： 1)森林土壤CH4吸收對N輸入響應機制的研究； 2)多種因素對林地土壤CH4吸收影響的長期定位試驗的研究； 3)甲烷氧化菌對各種干擾因素響應模式的研究； 4)氣候變化對林地土壤CH4吸收的影響及其響應機制的研究。

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(責任編輯 于靜嫻)

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。

浙江省自然科學基金項目(LY15C160004)； 浙江省科技創(chuàng)新團隊項目(2012R10030-11)。

S718.5

A

1001-7488(2017)05-0156-08

* 劉娟為通訊作者。