全球变化背景下森林凋落物分解的驱动动力学研究
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摘要
近年来,随着人类活动的日益频繁,尤其是城市化进程和工业化进程的快速推进,消耗了大量能源并释放出大量污染物,从而导致全球性的气候和环境变化,如高温、酸雨及大气氮沉降等。由于全球变化可能对森林生态系统造成多重影响等因素,全球变化已引起人们的广泛关注。但是,全球变化背景下森林凋落物分解的驱动动力学研究,尤其是全球性营养循环及其持续加速演变的全球变化对中国亚热带森林不同林型凋落物分解系统所造成的潜在影响,却知之甚少。
本研究选择中国南京紫金山地区中国亚热带森林两种林型的优势树种(麻栎Quercus acutissima及马尾松Pinus massoniana)的凋落物作为实验材料。两种凋落物与其原生土混合分别用高温(升温幅度：10℃,对照：20℃)、酸雨(低pH处理：4.5,高pH处理：5.0,对照：pH5.6)、不同沉降量的氮沉降(低量氮处理：1gNm-2yr-1,中量氮处理：2gNm-2yr-1,高量氮处理：3g N m-2yr-1,氮增施类型：NH4NO3,对照：水)及不同类型的氮沉降(氮增施类型：NH4+、NO3-、CO(NH2)2及前三者等比例配比的混合氮,氮增施量：3gNm-2yr-1,对照：水)进行处理。培养后测定凋落物的化学成分(木质素、全碳化合物及氮)、凋落物的失重速率、土壤pH值及其相关分解酶的活性。本研究旨在进行高温、酸雨、不同沉降量的氮沉降及不同类型的氮沉降背景下中国亚热带阔叶林(Quercus acutissima)及针叶林(Pinus massoniana)凋落物分解的驱动动力学的研究。本研究的主要目标是：(1)分析阔叶林凋落物及针叶林凋落物在高温、酸雨、不同沉降量的氮沉降及不同类型的氮沉降背景下分解的差异性及其敏感性；(2)探究高温、酸雨、不同沉降量的氮沉降及不同类型的氮沉降对中国亚热带森林凋落物分解及其相关分解酶活性的影响；(3)探析在高温、酸雨、不同沉降量的氮沉降及不同类型的氮沉降背景下驱动中国亚热带森林凋落物分解的主要分解酶。
本研究主要结果如下：
(1)温度升高(增幅为10℃)加速中国亚热带森林凋落物的分解速率,且阔叶林凋落物分解的温度敏感性与针叶林凋落物分解的温度敏感性无显著差异；高温增加微生物分解酶(过氧化氢酶、多酚氧化酶、纤维素酶、转化酶、尿酶、硝酸还原酶和酸性磷酸酶)的活性,且多酚氧化酶的温度敏感性显著大于其它微生物分解酶(过氧化氢酶、纤维素酶、转化酶、尿酶、硝酸还原酶和酸性磷酸酶)的温度敏感性；针叶林硝酸还原酶的温度敏感性显著大于阔叶林硝酸还原酶的温度敏感性,而其它微生物分解酶(过氧化氢酶、多酚氧化酶、纤维素酶、转化酶、尿酶和酸性磷酸酶)的温度敏感性在阔叶林和针叶林之间无显著差异；参与阔叶林凋落物分解的主要分解酶为过氧化氢酶和多酚氧化酶,而参与针叶林凋落物分解的主要分解酶为过氧化氢酶；高温促进阔叶林凋落物木质素的分解速率却减缓纤维素的分解速率,而高温促进针叶林凋落物纤维素的分解速率却减缓木质素的分解速率。根据本结果,长期的高温对森林凋落物分解的加速效应可能导致土壤碳库以二氧化碳的形式转移至大气中,从而改变森林生态系统的碳吸收及碳释放间的平衡,并进而加速全球气候变暖。
(2)酸雨抑制中国亚热带森林凋落物的分解速率,且酸雨对针叶林凋落物分解的抑制效应较大；酸雨降低微生物分解酶(多酚氧化酶、纤维素酶、转化酶、尿酶、硝酸还原酶、酸性磷酸酶和碱性磷酸酶)的活性,却增加过氧化氢酶的活性；在酸雨处理条件下,参与阔叶林凋落物分解的主要分解酶为过氧化氢酶和多酚氧化酶,而参与针叶林凋落物分解的主要分解酶为尿酶和硝酸还原酶；在对照处理条件下,参与阔叶林凋落物分解的主要分解酶为过氧化氢酶,而参与针叶林凋落物分解的主要分解酶为转化酶。根据本结果,长期的酸雨对森林凋落物分解的抑制效应可能导致土壤碳库的积累,从而改变生态系统碳流动、营养循环及腐殖质形成间的平衡,进而对森林生态系统造成多重影响。
(3)中量水平(2g N m-2yr-1)及高量水平(3g N m-2yr-1)的氮增施显著加速中国亚热带阔叶林凋落物的分解速率,而仅有高量水平(3g N m-2yr-1)的氮增施显著加速中国亚热带针叶林凋落物的分解速率,表明氮沉降条件下阔叶林凋落物分解的敏感性大于针叶林凋落物分解的敏感性；氮增施增加微生物分解酶(过氧化氢酶、多酚氧化酶、纤维素酶、转化酶、尿酶和酸性磷酸酶)的活性,却抑制硝酸还原酶的活性；在氮增施处理条件下,参与阔叶林凋落物分解的主要分解酶为过氧化氢酶、纤维素酶和酸性磷酸酶,而参与针叶林凋落物分解的主要分解酶为过氧化氢酶、纤维素酶、转化酶和硝酸还原酶；在对照处理条件下,参与阔叶林凋落物分解的主要分解酶为纤维素酶和硝酸还原酶,而参与针叶林凋落物分解的主要分解酶为过氧化氢酶、纤维素酶和酸性磷酸酶。根据本结果,长期的氮沉降对森林凋落物分解的增施效应可能增加大气中二氧化碳的相对含量,进而导致全球气候变暖。所以,长期的氮沉降可能引起多个气候及环境因子的相应变化,进而对森林生态系统造成多方面的影响。
(4)所有类型的氮增施(NH4+、N03-、CO(NH2)2及前三者等量配比的混合氮)均显著增加中国亚热带阔叶林凋落物的分解速率,而仅有混合氮及CO(NH2)2显著增加中国亚热带针叶林凋落物的分解速率,表明氮沉降条件下阔叶林凋落物分解的敏感性大于针叶林凋落物分解的敏感性；混合氮对两种凋落物微生物分解的增施效应均显著大于单一氮(NH4+,N03-及CO(NH2)2)的增施效应,有机氮(CO(NH2)2)对两种凋落物分解的增施效应均大于无机氮(NH4+及NO3-)的增施效应,而两种无机氮对两种凋落物分解的增施效应均无显著差异；所有类型的氮增施均增加微生物分解酶(过氧化氢酶、多酚氧化酶、纤维素酶、转化酶、尿酶、硝酸还原酶和酸性磷酸酶)的活性,且混合氮对分解酶活性的增施效应大于单一氮的增施效应；部分微生物分解酶(过氧化氢酶、硝酸还原酶及多酚氧化酶)的活性在有机氮增施条件下大于无机氮增施,而其它微生物分解酶(纤维素酶、转化酶、酸性磷酸酶及尿酶)的活性在无机氮增施条件下大于有机氮增施,表明参与凋落物分解的微生物对氮源的选择有些偏向于有机氮而有些偏向于无机氮。根据本结果,长期的氮沉降对凋落物分解的增施效应可能增加微生物对氮源的获取、植物对氮素的吸收及植物生物量的累积,进而增加微生物的代谢活性、森林凋落物的数量和质量,并加速森林凋落物的分解速率,从而导致森林生态系统对大气氮沉降响应的正反馈循环。
It is widely recognized that leaf litter decomposition and the accompanying release of nutrients and CO2and formation of SOM are fundamental processes in ecosystem nutrient cycling, C flux, and humus formation. Thus, the study of the processes and the drive kinetics of litter decomposition in subtropical forests in China is important to understanding these ecosystem functioning.
With the continuing increase in human activities in recent decades, such as urbanization and industrialization, which leading to a substantial increasing consumption of energy and thus emissions, there are induced climate and environment changes in global scale, such as the increased global temperature, the accelerated rates of acid rain and atmospheric N deposition. The global change may have multiple effects on forest ecosystems. Thus, their effects on ecological functioning of forest ecosystems, such as the drive kinetics of litter decomposition, have stimulated considerable interest, specifically in terms of the global nutrition cycle and its potential contribution to the rates of the ongoing global change in the coming decades.
Two dominant litter types were chosen from Zijin Mountain in China:Quercus acutissima leaves from a broad-leaved forest and Pinus massoniana needles from a coniferous forest. The litter samples were incubated in microcosms with their original forest soil and treated with high temperature (10℃warmer, and control:20℃), acid rain (low pH:4.5, high pH:5.0, control:pH5.6), gradient nitrogen deposition (low nitrogen:1g N m-2yr-1, medium nitrogen:2g N m-2yr-1, high nitrogen:3g N m-2yr-1, the form of nitrogen fertilization:NH4NO3, control:water), and different forms of N deposition (the form of nitrogen fertilization:NH4+, NO3-, CO(NH2)2, mixed N (a mix of all three equally), the level of nitrogen fertilization:3g N m-2yr-1, control: water), respectively. During the incubation, chemical composition (i.e., lignin, total carbohydrate, and nitrogen), litter mass losses, soil pH values, and the activities of degradative enzymes were determined. This study was carried out to assess the drive kinetics of litter decomposition under high temperature, acid rain, gradient N deposition, and different forms of N deposition in a subtropical broad-leaved forest (Quercus acutissima) and a coniferous forest (Pinus massoniana) in Zijin Mountain in China:(1) to determine the differences in litter decomposition between broadleaf forest leaves and coniferous forest needles in response to simulated high temperature, acid rain, gradient N deposition, and different forms of N deposition, respectively,(2) to assess the effects of simulated high temperature, acid rain, gradient N deposition, and different forms of N deposition on litter decomposition and related soil enzyme activities, respectively, and (3) to identify the major degradative enzyme contributors which drive litter decomposition under simulated high temperature, acid rain, gradient N deposition, and different forms of N deposition, respectively.
The results of this study are as follows:
(1) High temperature accelerates decomposition rates of the two litter types, and there was no significant difference in the temperature sensitivity of litter decomposition between broadleaf forest leaves and coniferous forest needles. High temperature also enhances soil enzyme activities (i.e., catalase, cellulase, invertase, polyphenol oxidase, nitrate reductase, urease, and acid phosphatase) in the two forest types, and the temperature sensitivity of polyphenol oxidase was significantly higher than those of the other soil enzymes (i.e., catalase, cellulase, invertase, nitrate reductase, urease, and acid phosphatase). Meanwhile, the temperature sensitivity of nitrate reductase was significantly higher in the coniferous forest soil than in the broadleaf forest soil, while there was no significant difference in the temperature sensitivity of the other soil enzymes (i.e., catalase, cellulase, invertase, polyphenol oxidase, urease, and acid phosphatase) between broadleaf forest and coniferous forest. Catalase and polyphenol oxidase were primarily responsible for litter decomposition in the broad-leaved forest, while catalase was primarily responsible for litter decomposition in the coniferous forest. High temperature tended to accelerate lignin decomposition rates but restrain cellulose decomposition rates for broadleaf forest leaves, while the opposite results were found for coniferous forest needles. The high temperature-accelerated litter decomposition rates may be partly attributed to the accelerated soil enzyme activities. As a long-term consequence, the high temperature-induced acceleration of litter decomposition rates in forest ecosystems may induce C stored belowground to be transferred to the atmosphere as greenhouse gas CO2, which may alter the balance between C uptake and release, and then alter the global C cycle.
(2) Acid rain reduces litter decomposition rates of the two little types, and the effects of acid rain on litter decomposition rates of coniferous forest needles were higher than on those of broadleaf forest leaves. Simulated acid rain also reduces the activities of cellulase, invertase, nitrate reductase, acid phosphatase, alkaline phosphatase, polyphenol oxidase, and urease, while it enhances catalase activities in most cases during litter decomposition. Catalase and polyphenol oxidase were primarily responsible for litter decomposition in acid treatments, and catalase was primarily responsible for litter decomposition in control treatment in the broad-leaved forest. Nitrate reductase and urease were primarily responsible for litter decomposition in acid treatments, and invertase was primarily responsible for litter decomposition in control treatment in the coniferous forest. According to the results of this study, soil C in subtropical forests would accumulate as a long-term consequence of continued acid rain. This may presumably alter the balance of ecosystem C flux, nutrients cycling, and humus formation, which may, in turn, have multiple effects on forest ecosystems.
(3) Medium-N and high-N fertilization significantly accelerates litter decomposition rates of broadleaf forest leaves, while only high-N fertilization significantly accelerates litter decomposition rates of coniferous forest needles, suggesting that the temperature sensitivities of litter decomposition for broadleaf forest leaves were significantly higher than that for coniferous forest needles under N deposition. N fertilization enhances the activities of catalase, cellulase, invertase, acid phosphatase, polyphenol oxidase, and urease, while it reduces nitrate reductase activities in most cases during litter decomposition. Cellulase and nitrate reductase were primarily responsible for litter decomposition in the broad-leaved forest, while catalase, cellulase, and acid phosphatase were primarily responsible for litter decomposition in the coniferous forest under no N fertilization; catalase, cellulase and acid phosphatase were primarily responsible for litter decomposition in the broad-leaved forest, while catalase, cellulase, invertase, and nitrate reductase were primarily responsible for litter decomposition in the coniferous forest under N fertilization. The accelerated litter decomposition rates under N fertilization may cause an increase of the greenhouse gas CO2in the atmosphere. Elevated atmospheric CO2concentration might also lead to a warmer climate, which may enhances the rates of litter decomposition in forest ecosystems. Thus, anthropogenic N deposition can lead to related changes in climatic factors, which, in turn, generally have multiple effects on forest ecosystems.
(4) All four forms of N fertilization [NH4+, NO3-, CO(NH2)2, and a mix of all three] significantly accelerates litter decomposition rates in the broadleaf forest, while only the mixed N and CO(NH2)2fertilization significantly accelerates litter decomposition rates in the coniferous forest, suggesting that the temperature sensitivities of litter decomposition for broadleaf forest leaves were significantly higher than that for coniferous forest needles under N deposition. Litter decomposition rates under the mixed N fertilization were significantly higher than those under any single form of N fertilization. All forms of N fertilization enhance soil enzyme activities (i.e., catalase, cellulase, invertase, polyphenol oxidase, nitrate reductase, urease, and acid phosphatase) during litter decomposition in the two forest types. Soil enzyme activities under the mixed N fertilization were higher than those under any single form of N fertilization. Catalase, nitrate reductase, and polyphenol oxidase activities under inorganic N fertilization were higher than those under organic N fertilization, while cellulase, invertase, acid phosphatase, and urease activities under organic N fertilization were higher than those under inorganic N fertilization, suggesting that some groups of soil microorganisms that secrete exoenzymes for litter decomposition may prefer either organic N or inorganic N. The accelerated litter decomposition rates under N fertilization may cause an increase in N availability for micro-decomposer metabolism, greater N uptake for plants, and possibly a greater amount of C sequestered in plant biomass as a long-term consequence of continued N deposition, following this, the activities of micro-decomposer metabolism, the amount of litter fall and the quality of litter may all increase, which, in turn, could enhances litter decomposition rates in forest ecosystems and initiate a so-called feed-forward cycle.
本研究选择中国南京紫金山地区中国亚热带森林两种林型的优势树种(麻栎Quercus acutissima及马尾松Pinus massoniana)的凋落物作为实验材料。两种凋落物与其原生土混合分别用高温(升温幅度：10℃,对照：20℃)、酸雨(低pH处理：4.5,高pH处理：5.0,对照：pH5.6)、不同沉降量的氮沉降(低量氮处理：1gNm-2yr-1,中量氮处理：2gNm-2yr-1,高量氮处理：3g N m-2yr-1,氮增施类型：NH4NO3,对照：水)及不同类型的氮沉降(氮增施类型：NH4+、NO3-、CO(NH2)2及前三者等比例配比的混合氮,氮增施量：3gNm-2yr-1,对照：水)进行处理。培养后测定凋落物的化学成分(木质素、全碳化合物及氮)、凋落物的失重速率、土壤pH值及其相关分解酶的活性。本研究旨在进行高温、酸雨、不同沉降量的氮沉降及不同类型的氮沉降背景下中国亚热带阔叶林(Quercus acutissima)及针叶林(Pinus massoniana)凋落物分解的驱动动力学的研究。本研究的主要目标是：(1)分析阔叶林凋落物及针叶林凋落物在高温、酸雨、不同沉降量的氮沉降及不同类型的氮沉降背景下分解的差异性及其敏感性；(2)探究高温、酸雨、不同沉降量的氮沉降及不同类型的氮沉降对中国亚热带森林凋落物分解及其相关分解酶活性的影响；(3)探析在高温、酸雨、不同沉降量的氮沉降及不同类型的氮沉降背景下驱动中国亚热带森林凋落物分解的主要分解酶。
本研究主要结果如下：
(1)温度升高(增幅为10℃)加速中国亚热带森林凋落物的分解速率,且阔叶林凋落物分解的温度敏感性与针叶林凋落物分解的温度敏感性无显著差异；高温增加微生物分解酶(过氧化氢酶、多酚氧化酶、纤维素酶、转化酶、尿酶、硝酸还原酶和酸性磷酸酶)的活性,且多酚氧化酶的温度敏感性显著大于其它微生物分解酶(过氧化氢酶、纤维素酶、转化酶、尿酶、硝酸还原酶和酸性磷酸酶)的温度敏感性；针叶林硝酸还原酶的温度敏感性显著大于阔叶林硝酸还原酶的温度敏感性,而其它微生物分解酶(过氧化氢酶、多酚氧化酶、纤维素酶、转化酶、尿酶和酸性磷酸酶)的温度敏感性在阔叶林和针叶林之间无显著差异；参与阔叶林凋落物分解的主要分解酶为过氧化氢酶和多酚氧化酶,而参与针叶林凋落物分解的主要分解酶为过氧化氢酶；高温促进阔叶林凋落物木质素的分解速率却减缓纤维素的分解速率,而高温促进针叶林凋落物纤维素的分解速率却减缓木质素的分解速率。根据本结果,长期的高温对森林凋落物分解的加速效应可能导致土壤碳库以二氧化碳的形式转移至大气中,从而改变森林生态系统的碳吸收及碳释放间的平衡,并进而加速全球气候变暖。
(2)酸雨抑制中国亚热带森林凋落物的分解速率,且酸雨对针叶林凋落物分解的抑制效应较大；酸雨降低微生物分解酶(多酚氧化酶、纤维素酶、转化酶、尿酶、硝酸还原酶、酸性磷酸酶和碱性磷酸酶)的活性,却增加过氧化氢酶的活性；在酸雨处理条件下,参与阔叶林凋落物分解的主要分解酶为过氧化氢酶和多酚氧化酶,而参与针叶林凋落物分解的主要分解酶为尿酶和硝酸还原酶；在对照处理条件下,参与阔叶林凋落物分解的主要分解酶为过氧化氢酶,而参与针叶林凋落物分解的主要分解酶为转化酶。根据本结果,长期的酸雨对森林凋落物分解的抑制效应可能导致土壤碳库的积累,从而改变生态系统碳流动、营养循环及腐殖质形成间的平衡,进而对森林生态系统造成多重影响。
(3)中量水平(2g N m-2yr-1)及高量水平(3g N m-2yr-1)的氮增施显著加速中国亚热带阔叶林凋落物的分解速率,而仅有高量水平(3g N m-2yr-1)的氮增施显著加速中国亚热带针叶林凋落物的分解速率,表明氮沉降条件下阔叶林凋落物分解的敏感性大于针叶林凋落物分解的敏感性；氮增施增加微生物分解酶(过氧化氢酶、多酚氧化酶、纤维素酶、转化酶、尿酶和酸性磷酸酶)的活性,却抑制硝酸还原酶的活性；在氮增施处理条件下,参与阔叶林凋落物分解的主要分解酶为过氧化氢酶、纤维素酶和酸性磷酸酶,而参与针叶林凋落物分解的主要分解酶为过氧化氢酶、纤维素酶、转化酶和硝酸还原酶；在对照处理条件下,参与阔叶林凋落物分解的主要分解酶为纤维素酶和硝酸还原酶,而参与针叶林凋落物分解的主要分解酶为过氧化氢酶、纤维素酶和酸性磷酸酶。根据本结果,长期的氮沉降对森林凋落物分解的增施效应可能增加大气中二氧化碳的相对含量,进而导致全球气候变暖。所以,长期的氮沉降可能引起多个气候及环境因子的相应变化,进而对森林生态系统造成多方面的影响。
(4)所有类型的氮增施(NH4+、N03-、CO(NH2)2及前三者等量配比的混合氮)均显著增加中国亚热带阔叶林凋落物的分解速率,而仅有混合氮及CO(NH2)2显著增加中国亚热带针叶林凋落物的分解速率,表明氮沉降条件下阔叶林凋落物分解的敏感性大于针叶林凋落物分解的敏感性；混合氮对两种凋落物微生物分解的增施效应均显著大于单一氮(NH4+,N03-及CO(NH2)2)的增施效应,有机氮(CO(NH2)2)对两种凋落物分解的增施效应均大于无机氮(NH4+及NO3-)的增施效应,而两种无机氮对两种凋落物分解的增施效应均无显著差异；所有类型的氮增施均增加微生物分解酶(过氧化氢酶、多酚氧化酶、纤维素酶、转化酶、尿酶、硝酸还原酶和酸性磷酸酶)的活性,且混合氮对分解酶活性的增施效应大于单一氮的增施效应；部分微生物分解酶(过氧化氢酶、硝酸还原酶及多酚氧化酶)的活性在有机氮增施条件下大于无机氮增施,而其它微生物分解酶(纤维素酶、转化酶、酸性磷酸酶及尿酶)的活性在无机氮增施条件下大于有机氮增施,表明参与凋落物分解的微生物对氮源的选择有些偏向于有机氮而有些偏向于无机氮。根据本结果,长期的氮沉降对凋落物分解的增施效应可能增加微生物对氮源的获取、植物对氮素的吸收及植物生物量的累积,进而增加微生物的代谢活性、森林凋落物的数量和质量,并加速森林凋落物的分解速率,从而导致森林生态系统对大气氮沉降响应的正反馈循环。
It is widely recognized that leaf litter decomposition and the accompanying release of nutrients and CO2and formation of SOM are fundamental processes in ecosystem nutrient cycling, C flux, and humus formation. Thus, the study of the processes and the drive kinetics of litter decomposition in subtropical forests in China is important to understanding these ecosystem functioning.
With the continuing increase in human activities in recent decades, such as urbanization and industrialization, which leading to a substantial increasing consumption of energy and thus emissions, there are induced climate and environment changes in global scale, such as the increased global temperature, the accelerated rates of acid rain and atmospheric N deposition. The global change may have multiple effects on forest ecosystems. Thus, their effects on ecological functioning of forest ecosystems, such as the drive kinetics of litter decomposition, have stimulated considerable interest, specifically in terms of the global nutrition cycle and its potential contribution to the rates of the ongoing global change in the coming decades.
Two dominant litter types were chosen from Zijin Mountain in China:Quercus acutissima leaves from a broad-leaved forest and Pinus massoniana needles from a coniferous forest. The litter samples were incubated in microcosms with their original forest soil and treated with high temperature (10℃warmer, and control:20℃), acid rain (low pH:4.5, high pH:5.0, control:pH5.6), gradient nitrogen deposition (low nitrogen:1g N m-2yr-1, medium nitrogen:2g N m-2yr-1, high nitrogen:3g N m-2yr-1, the form of nitrogen fertilization:NH4NO3, control:water), and different forms of N deposition (the form of nitrogen fertilization:NH4+, NO3-, CO(NH2)2, mixed N (a mix of all three equally), the level of nitrogen fertilization:3g N m-2yr-1, control: water), respectively. During the incubation, chemical composition (i.e., lignin, total carbohydrate, and nitrogen), litter mass losses, soil pH values, and the activities of degradative enzymes were determined. This study was carried out to assess the drive kinetics of litter decomposition under high temperature, acid rain, gradient N deposition, and different forms of N deposition in a subtropical broad-leaved forest (Quercus acutissima) and a coniferous forest (Pinus massoniana) in Zijin Mountain in China:(1) to determine the differences in litter decomposition between broadleaf forest leaves and coniferous forest needles in response to simulated high temperature, acid rain, gradient N deposition, and different forms of N deposition, respectively,(2) to assess the effects of simulated high temperature, acid rain, gradient N deposition, and different forms of N deposition on litter decomposition and related soil enzyme activities, respectively, and (3) to identify the major degradative enzyme contributors which drive litter decomposition under simulated high temperature, acid rain, gradient N deposition, and different forms of N deposition, respectively.
The results of this study are as follows:
(1) High temperature accelerates decomposition rates of the two litter types, and there was no significant difference in the temperature sensitivity of litter decomposition between broadleaf forest leaves and coniferous forest needles. High temperature also enhances soil enzyme activities (i.e., catalase, cellulase, invertase, polyphenol oxidase, nitrate reductase, urease, and acid phosphatase) in the two forest types, and the temperature sensitivity of polyphenol oxidase was significantly higher than those of the other soil enzymes (i.e., catalase, cellulase, invertase, nitrate reductase, urease, and acid phosphatase). Meanwhile, the temperature sensitivity of nitrate reductase was significantly higher in the coniferous forest soil than in the broadleaf forest soil, while there was no significant difference in the temperature sensitivity of the other soil enzymes (i.e., catalase, cellulase, invertase, polyphenol oxidase, urease, and acid phosphatase) between broadleaf forest and coniferous forest. Catalase and polyphenol oxidase were primarily responsible for litter decomposition in the broad-leaved forest, while catalase was primarily responsible for litter decomposition in the coniferous forest. High temperature tended to accelerate lignin decomposition rates but restrain cellulose decomposition rates for broadleaf forest leaves, while the opposite results were found for coniferous forest needles. The high temperature-accelerated litter decomposition rates may be partly attributed to the accelerated soil enzyme activities. As a long-term consequence, the high temperature-induced acceleration of litter decomposition rates in forest ecosystems may induce C stored belowground to be transferred to the atmosphere as greenhouse gas CO2, which may alter the balance between C uptake and release, and then alter the global C cycle.
(2) Acid rain reduces litter decomposition rates of the two little types, and the effects of acid rain on litter decomposition rates of coniferous forest needles were higher than on those of broadleaf forest leaves. Simulated acid rain also reduces the activities of cellulase, invertase, nitrate reductase, acid phosphatase, alkaline phosphatase, polyphenol oxidase, and urease, while it enhances catalase activities in most cases during litter decomposition. Catalase and polyphenol oxidase were primarily responsible for litter decomposition in acid treatments, and catalase was primarily responsible for litter decomposition in control treatment in the broad-leaved forest. Nitrate reductase and urease were primarily responsible for litter decomposition in acid treatments, and invertase was primarily responsible for litter decomposition in control treatment in the coniferous forest. According to the results of this study, soil C in subtropical forests would accumulate as a long-term consequence of continued acid rain. This may presumably alter the balance of ecosystem C flux, nutrients cycling, and humus formation, which may, in turn, have multiple effects on forest ecosystems.
(3) Medium-N and high-N fertilization significantly accelerates litter decomposition rates of broadleaf forest leaves, while only high-N fertilization significantly accelerates litter decomposition rates of coniferous forest needles, suggesting that the temperature sensitivities of litter decomposition for broadleaf forest leaves were significantly higher than that for coniferous forest needles under N deposition. N fertilization enhances the activities of catalase, cellulase, invertase, acid phosphatase, polyphenol oxidase, and urease, while it reduces nitrate reductase activities in most cases during litter decomposition. Cellulase and nitrate reductase were primarily responsible for litter decomposition in the broad-leaved forest, while catalase, cellulase, and acid phosphatase were primarily responsible for litter decomposition in the coniferous forest under no N fertilization; catalase, cellulase and acid phosphatase were primarily responsible for litter decomposition in the broad-leaved forest, while catalase, cellulase, invertase, and nitrate reductase were primarily responsible for litter decomposition in the coniferous forest under N fertilization. The accelerated litter decomposition rates under N fertilization may cause an increase of the greenhouse gas CO2in the atmosphere. Elevated atmospheric CO2concentration might also lead to a warmer climate, which may enhances the rates of litter decomposition in forest ecosystems. Thus, anthropogenic N deposition can lead to related changes in climatic factors, which, in turn, generally have multiple effects on forest ecosystems.
(4) All four forms of N fertilization [NH4+, NO3-, CO(NH2)2, and a mix of all three] significantly accelerates litter decomposition rates in the broadleaf forest, while only the mixed N and CO(NH2)2fertilization significantly accelerates litter decomposition rates in the coniferous forest, suggesting that the temperature sensitivities of litter decomposition for broadleaf forest leaves were significantly higher than that for coniferous forest needles under N deposition. Litter decomposition rates under the mixed N fertilization were significantly higher than those under any single form of N fertilization. All forms of N fertilization enhance soil enzyme activities (i.e., catalase, cellulase, invertase, polyphenol oxidase, nitrate reductase, urease, and acid phosphatase) during litter decomposition in the two forest types. Soil enzyme activities under the mixed N fertilization were higher than those under any single form of N fertilization. Catalase, nitrate reductase, and polyphenol oxidase activities under inorganic N fertilization were higher than those under organic N fertilization, while cellulase, invertase, acid phosphatase, and urease activities under organic N fertilization were higher than those under inorganic N fertilization, suggesting that some groups of soil microorganisms that secrete exoenzymes for litter decomposition may prefer either organic N or inorganic N. The accelerated litter decomposition rates under N fertilization may cause an increase in N availability for micro-decomposer metabolism, greater N uptake for plants, and possibly a greater amount of C sequestered in plant biomass as a long-term consequence of continued N deposition, following this, the activities of micro-decomposer metabolism, the amount of litter fall and the quality of litter may all increase, which, in turn, could enhances litter decomposition rates in forest ecosystems and initiate a so-called feed-forward cycle.
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