亚高山森林植物层次去除对土壤生态过程的影响
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摘要
生物多样性与生态系统功能的关系是生态学领域研究的热点与难点,但现有的研究主要集中在草地生态系统、水生生态系统和农田生态系统,受研究手段的限制,对森林生态系统研究相对较少。川西亚高山针叶林位于我国青藏高原东缘和长江上游地区,在生物多样性保育、水源涵养和全球碳循环中具有十分重要的、不可替代的作用和地位。同时,川西亚高山针叶林具有层次结构和物种丰富度相对简单、更新和演替规律十分清晰、结构和功能类型更加分化等特点,这为研究森林物种组成和多样性变化对森林生态系统功能及过程的影响提供了理想的天然实验室。大量研究表明,土壤生物多样性、植被与生态系统过程密切相关,土壤生物群落与植被的互动决定着生态系统过程,土壤生态系统很可能就是联系生物多样性与生态系统功能关系的桥梁与纽带。基于此,本研究于2007-2009年,以1988年营造的云杉人工纯林与混交林为研究对象,运用野外定位观测和林地控制实验的手段,从“植物多样性对土壤动物的影响、对土壤微生物的影响、对土壤酶活性的影响、对凋落物分解的影响”等4个方面,探讨了植物多样性对土壤生态过程的影响。以期丰富森林生态系统生物多样性与生态系统功能的理论研究成果,并为川西亚高山退化森林生态系统的恢复重建提供必要的基础数据。主要研究结果如下：
1.植物多样性对土壤动物的影响
不同林分类型的土壤动物群落结构具有明显的季节变化。云杉纯林获土壤动物7197头(不包括原生动物,下同)；云杉混交林获土壤动物8831头。云杉混交林各类土壤动物类群数、密度、生物量和多样性指数均高于纯林,其中,大型土壤动物林分间有显著或极显著差异。云杉混交林腐食性和捕食性土壤动物比例高丁。纯林,而植食性土壤动物比例低于纯林。A/C值受纬度和海拔的影响,一定程度上能反映不同地带土壤动物的分布特征；在同一地带,还与生境的异质性、受干扰的程度和植物群落的多样性有关。云杉纯林A/C值为0.87,混交林为1.59,约为纯林1.83倍。云杉纯林土壤原生动物密度为365头／克鲜土,混交林为1046头／克鲜士,极显著高于纯林。两类林分土壤动物的剖面分布均具明显表聚性,并且有相似的季节动态。
除草处理(RH)获土壤动物7529头；除灌处理(RS)获土壤动物6103头；对照(CK)获土壤动物10986头。除灌除草处理后,大型土壤动物类群数短期内有增加的趋势,而中小型土壤动物类群数和土壤动物总类群数呈下降趋势；除灌除草处理后,各类：L壤动物密度、生物量和DG指数基本以CK>RH>RS；腐食性土壤动物数量和比例以及A/C值均有所下降,其中,RH处理A/C值为0.92,RS处理A/C值为0.82,CK处理A/C值为1.43;除灌除草处理后,土壤原生动物密度下降,RH处理为958头／克鲜土,RS处理为1510头／克鲜土,CK处理为2666头／克鲜土,处理间有极显著差异；除灌除草处理后,土壤动物剖面分布的表聚性没有改变,但递减的速率以RS>RH>CK。总体而言,除灌处理对土壤动物群落的影响大于除草处理。
2.植物多样性对土壤微生物的影响
不同林分类型的土壤微生物群落的组成、数量、生物量存在差异。云杉纯林与混交林虽均以细菌数量最多,放线菌次之,真菌最少,但混交林真菌比例大于纯林;云杉纯林细菌数量和土壤微生物总数显著低于混交林,而放线菌和真菌数量极显著低于混交林；两类林分细菌、真菌、土壤微生物总数及纯林放线菌的剖面分布具明显表聚性,而混交林放线菌则以凋落物层最低,0-5cm土层最高；云杉纯林与混交林各类土壤微生物数量均以夏季最高,春季次之,秋季最低；土壤微生物生物量是土壤有机质较易分解的部分,是土壤养分重要的源与库,云杉纯林土壤微生物生物量碳氮低于混交林,其剖面分布和季节动态与土壤可培养微生物数量一致。
除灌除草处理对土壤微生物数量和生物量的剖面分布特征产生了不同程度的影响。土壤细菌和真菌数量均以CK最大,RH次之,RS最小,而上壤放线菌数量则以RH最大,CK次之,RS最小；除灌除草处理对不同层次土壤微生物生物量的影响不同,其中,凋落物层土壤微生物生物量以RH最高,CK次之,RS最小,而0-5cm、5-10cm、10-15cm土层则以CK最高,RH次之,RS最低,除灌处理对土壤微生物生物量的影响大于除草处理,除草处理后土壤微生物生物量短期内有升高的趋势；除灌除草处理后,土壤微生物的剖面分布格局没有发生太大变化,但随土层加深,微生物数量和生物量递减的速率表现出差异,以RS最大,CK最小。
3.植物多样性对上壤酶活性的影响
不同林分类型的土壤蔗糖酶、脲酶、纤维素酶和脱氢酶活性表现出一定差异。云杉混交林各种酶活性均高于纯林；两类林分土壤酶活性均具明显表聚性,且有相似的季节动态,其中,蔗糖酶、脲酶与脱氢酶活性以秋季最低,夏季或春季最高,纤维素酶活性以夏季或秋季最高,春季最低。
除灌除草处理后,4种酶活性均降低,以CK>RH>RS,除灌处理的影响大于除草处理；4种酶活性均以CK随土层加深递减的速率最小,RS递减的速率最大；4种酶中,以纤维素酶活性随土层加深递减的速率最大,脱氢酶活性递减的速率最小。
4.植物多样性对凋落物分解的影响
不同林分类型的凋落物分解特征存在差异。云杉纯林凋落物分解速率低于混交林,约为混交林的65.51-99.63%,前期差异明显,后期差异较小。云杉纯林土壤微生物与小型土壤动物(主要是线虫和原生动物)对凋落物分解的贡献率达68.71-94.03%,大中型土壤动物对凋落物分解的贡献率为5.97～31.29%；云杉混交林土壤微生物与小型土壤动物对凋落物分解的贡献率达90.29～94.22%,大中型土壤动物对凋落物分解的贡献率为5.78-9.71%。各类土壤动物与微生物的共同作用大于仅微生物和小型湿生上壤动物的作用。在云杉凋落物分解的开始2年,除N元素出现富集外,P、K、Ca、Mg等元素均表现为释放,从释放量的大小来看,基本以混交林大于纯林。在该研究区域,所有分解者中以线虫、原生动物和真菌对凋落物分解起至关重要的作用。
除灌除草处理降低了凋落物分解速率,并且除灌处理导致的下降程度高于除草处理；各处理土壤动物与微生物对凋落物分解的共同作用大于仅微生物和小型湿生土壤动物的作用。各处理云杉凋落物分解的开始2年,N元素表现为富集,而P、K、Ca、Mg等元素则表现为释放。其中,除灌处理对凋落物养分释放表现为不利的影响,除Mg元素以外的N、P、K、Ca元素均以除灌处理释放量最小；除草处理在一定程度上有利于K元素的释放,而对N、P、Ca元素的释放影响不大,释放量仅次于或略等于对照。除灌除草处理后,凋落物分解过程中土壤动物、微生物及酶活性的变化并不是简单的以CK>RH>RS,而是表现出更加复杂的模式。由此认为,灌草层去除引起的凋落物组成和质量的改变,以及凋落物分解过程中土壤生物的多样性格局及酶活性的变化趋势是影响凋落物分解和养分周转的重要因素；同时,去除行为本身可能对森林生态系统的生物多样性以及物质循环功能产生干扰,干扰效应在处理早期比较明显,随时间推移逐渐减弱。
综上所述,植物多样性低的林分,其土壤生物多样性、土壤酶活性以及凋落物分解速率低于多样性高的林分；灌木层缺失对土壤生态过程的影响大于草本层缺失。因此,提高植物多样性,丰富凋落物组成,通过地上／地下生物互动,促进养分元素的循环,是增强生态系统各项功能的有效途径。
The relationship between biodiversity and ecosystem function is a hot and difficult problem in the field of ecological research, but a lot of studies are mainly concentrated on the research of grassland ecosystems, aquatic ecosystems and farmland ecosystem, because the means of research is inadequate, studies of forest ecosystem are relatively seldom, as compared with those ecosystems. Subalpine coniferous forests located in eastern edge of Qinghai-Tibet Plateau and the upper area of Yangtze River, it played an important and irreplaceable role in biodiversity conservation, water conservation and the global carbon cycle. Meanwhile, subalpine coniferous forests have some characteristics, for example, the hierarchical structure is clear and species richness is relatively simple, the law of update and succession is clearer; the type of structure and function is more various and so on. Those characteristics provide an ideal natural laboratory for the study of the composition of forest species and variations of forest ecosystems function and the process as affected by diversity change. A large number of studies have shown that the process is closely related to soil biodiversity, vegetation and ecosystems, interaction relationship of soil biological communities and vegetation determine ecosystem processes, the soil ecosystem is likely a bridge which makes biodiversity contact with ecosystem function. Above all, spruce artificial pure forest and mixed forest which were planted in1988were studied in this paper, the effects of plant diversity on soil microbes, soil enzyme activity, litter decomposition though field positioning observation and woodland means of control experiments and the influence of plant diversity on soil ecological process. Those results would enrich theoretical studies of the forest ecosystem biodiversity and ecosystem function, and provide basic data for restoration and reconstruction of subalpine degraded forest ecosystems. The main findings are as follows:
1. Effects of plant diversity on soil fauna
Different types of stand showed differences of the structure and spatial and temporal distribution of soil animal community. There are7197soil animals in Picea asperata pure plantation(not including protozoa, the same below); there are8831soil animals in Picea asperata mixed plantation. The number, density, biomass and diversity index of soil animal group of Picea asperata mixed plantation were higher than Picea asperata pure plantation. Among them, the number, density, biomass and diversity indices of macro-fauna have a significant or very significant difference. The quantities of saprophagous and predatory soil fauna in Picea asperata mixed plantation are higher than Picea asperata pure plantation, but the quantities of phytophagous soil animals are less than Picea asperata pure plantation. The ratio of A/C was effected by latitude and altitude, to some extent, reflect the characteristics of the distribution of soil animals in different zones; it is related to the heterogeneity of habitat, the degree of interference and diversity of plant communities in the same zone. The ratio of A/C of Picea asperata pure plantation was0.87, Picea asperata mixed plantation wasl.59and is about1.83times former. Soil protozoa density of Picea asperata pure plantation was365in1g fresh soil, soil protozoa density of spruce mixed forest was1046in1g fresh soil was significantly higher than Picea asperata pure plantation. The profile distribution of soil fauna of two stand types both have apparent surface gathering, and similar seasonal dynamics, are higher in summer than in spring and autumn.
Structure and distribution of soil animal communities showed different characteristics among different treatments.7529soil animals were collected in herb removal treatment (RH); the quantities of soil animals were6103in shrub removal treatment (RS).10986soil animals were collected in CK treatment. Through RH and RS treatment, the groups of soil macro-fauna showed increasing trend in short term, but the number of micro-and meso-soil fauna were decreasing; through RH and RS treatment, the density, biomass, and DG index of soil animals showed CK>RH>RS; number and proportion of saprophagous soil animals as well as A/C ratio were decreased, value of A/C was0.92in the RH treatment, value of A/C was0.82in the RS treatment, value of A/C of CK treatment was1.43; Through RH and RS treatment, soil protozoa density decreased,there were958soil animals in1g of fresh soil in RH treatment, there were1510soil animals in1g of fresh soil in RS treatment, there were2666soil animals in CK treatment, there was a significant difference among treatments; through RH and RS treatment, the profile distribution of soil animals has not changed, but a decreasing rate showed RS>RH>CK. Overall, effects of RS treatment on soil animal communities were more than RH treatment.
2. The effects of plant diversity on soil microbes
There were different component, number, biomass and temporal and spatial distribution of soil microbes in different plant types. Bacteria had the largest number in Picea asperata pure plantation and Picea asperata mixed plantation compared with other microbes, the order was Bacteria>actinomycetes>fungi, but the fungi ratio in mixed plantation was higher than that in pure plantation. The total number of microbes in Picea asperata pure plantation was significantly lower than that in Picea asperata mixed plantation, and number of actinomycetes and fungi in pure plantation were extremely lower than that in mixed plantation. Bacteria, fungi, total soil microbe number and actinomycetes in pure plantation in surface soil layer was higher than that in other soil layers, the actinomycetes had the lowest number in litter layer, and highest number in0-5cm layer compared with other soil layers. The order of Biomass in Picea asperata pure and mixed plantation was summer>spring>autumn. Biomass of soil microbes is the easy to decompose part in soil organic matter, which is an important source and sink, the profile distribution and seasonal dynamics of C/N of microbe biomass had the same trends with microbes number, and Picea asperata mixed plantation higher than that in Picea asperata pure plantation.
After shrub removal, soil microbes number, biomass and profile distribution changed a lot. The number order of soil bacteria and fungi was CK>RH>RS, and the number order of actinomycetes was RH>CK>RS. There was different affection on the microbes'biomass of soil layers after shrub and herb removal. The order of microbes'biomass in litter layer was RH>CK>RS, and The order of microbes'biomass in soil layers were CK>RH>RS, microbes' biomass had negative response after shrub removal, and had positive response after herb removal after a short period. There no huge changes in microbe profile distribution after shrub and herb removal, but microbe number and biomass reduced with the soil depth increased, the highest was RS, and the lowest was CK.
3. The effects of plant diversity on soil soil enzyme activity
There were diffrent activity of soil invertase, urease, cellulase and dehydrogenase in different plant types. All enzyme activity in the Picea asperata mixed plantation was higher than that in pure plantation. And enzyme activity in surface soil layer was higher than that in other soil layers, and had the same seasonal trends. The order of invertase, urease and dehydrogenase activity was autumnspring.
After shrub cutting,4enzymes'activity reduced, the order was CK>RH>RS, and the enzymes'activity effected by removing shrub are higher than that of removing herb. The lowest degressive rate of4enzymes activity with the soil depth increased was CK, the highest was RS. And cellulase had the highest degressive rate with the soil depth increased among the4enzymes, and dehydrogenase had the lowest degressive rate.
4. The effects of plant diversity on litter decomposition
Litter decomposition had different characteristics in the different plantation. The decomposition rate of Picea asperata pure plantation was lower than that in the Picea asperata mixed plantation, decomposition ratio of Picea asperata pure plantation and mixed plantation is65.51～99.63%. the significant difference was found in early period, and there was tiny difference in later period. The contribution of soil microbes and meso-fauna (mainly nematode and protozoa) to litter decomposition in Picea asperata pure plantation are68.71～94.03%, contribution of soil macro-fauna to litter decomposition was5.97～31.29%; The contribution of soil microbes and meso-fauna to litter decomposition in Picea asperata mixed plantation are90.29～94.22%, contribution of soil macro and meso-fauna to litter decomposition was5.78～9.71%. The interaction decomposition of soil microbes and soil fauna were higher that effect of pure soil microbes or soil fauna. The front2years' decomposition, P、K、Ca、Mg element showed releasing, but the N showed enrichment. Releasing quantity of mixed plantation was higher that in pure plantation.
After shrub and herb removal, the litter decomposition rate decreased, and litter decomposition rate decreased caused by shrub removal was higher than that in herb removal. The interaction decomposition of soil microbes and soil fauna were higher that effect of pure soil microbes or soil fauna. The front2years' decomposition, P、K、Ca、Mg element showed releasing, but the N showed enrichment. Shrub removal had negative effect on the litter decomposition, N、P、K、Ca element showed releasing excepting Mg. Herb removal had positive effect on K releasing, and had little affection on the N、P、Ca releasing, the releasing quantity had no significant difference compared with CK. After shrub and herb removal, the changes of soil fauna, soil microbes and enzyme activity during the litter decomposition hadn't the order CK>RH>RS, but had more complicated models. The result can be conclude that the shrub and herb removal lead to the composition and quality changes, and the soil fauna diversity and the change trends of enzyme were the main factors which lead to litter decomposition and nutrition cycle during the litter decomposition. Meanwhile, the removal is an important interference which affects the biodiversity and material cycle; the interference affection was significant in early period.
To summary, compared with rich plant diversity plantation, the soil biodiversity, soil enzyme activity, and litter decomposition rate were lower than that in the poor plant diversity plantation. Stronger affection on the soil ecology process was found in shrub removal compared with herb removal. Therefore, to increase plant diversity, to enrich litter composition, to interact upper and underground organisms, to improve nutrition recycles, can enhanced the functions of forest ecosystem.
1.植物多样性对土壤动物的影响
不同林分类型的土壤动物群落结构具有明显的季节变化。云杉纯林获土壤动物7197头(不包括原生动物,下同)；云杉混交林获土壤动物8831头。云杉混交林各类土壤动物类群数、密度、生物量和多样性指数均高于纯林,其中,大型土壤动物林分间有显著或极显著差异。云杉混交林腐食性和捕食性土壤动物比例高丁。纯林,而植食性土壤动物比例低于纯林。A/C值受纬度和海拔的影响,一定程度上能反映不同地带土壤动物的分布特征；在同一地带,还与生境的异质性、受干扰的程度和植物群落的多样性有关。云杉纯林A/C值为0.87,混交林为1.59,约为纯林1.83倍。云杉纯林土壤原生动物密度为365头／克鲜土,混交林为1046头／克鲜士,极显著高于纯林。两类林分土壤动物的剖面分布均具明显表聚性,并且有相似的季节动态。
除草处理(RH)获土壤动物7529头；除灌处理(RS)获土壤动物6103头；对照(CK)获土壤动物10986头。除灌除草处理后,大型土壤动物类群数短期内有增加的趋势,而中小型土壤动物类群数和土壤动物总类群数呈下降趋势；除灌除草处理后,各类：L壤动物密度、生物量和DG指数基本以CK>RH>RS；腐食性土壤动物数量和比例以及A/C值均有所下降,其中,RH处理A/C值为0.92,RS处理A/C值为0.82,CK处理A/C值为1.43;除灌除草处理后,土壤原生动物密度下降,RH处理为958头／克鲜土,RS处理为1510头／克鲜土,CK处理为2666头／克鲜土,处理间有极显著差异；除灌除草处理后,土壤动物剖面分布的表聚性没有改变,但递减的速率以RS>RH>CK。总体而言,除灌处理对土壤动物群落的影响大于除草处理。
2.植物多样性对土壤微生物的影响
不同林分类型的土壤微生物群落的组成、数量、生物量存在差异。云杉纯林与混交林虽均以细菌数量最多,放线菌次之,真菌最少,但混交林真菌比例大于纯林;云杉纯林细菌数量和土壤微生物总数显著低于混交林,而放线菌和真菌数量极显著低于混交林；两类林分细菌、真菌、土壤微生物总数及纯林放线菌的剖面分布具明显表聚性,而混交林放线菌则以凋落物层最低,0-5cm土层最高；云杉纯林与混交林各类土壤微生物数量均以夏季最高,春季次之,秋季最低；土壤微生物生物量是土壤有机质较易分解的部分,是土壤养分重要的源与库,云杉纯林土壤微生物生物量碳氮低于混交林,其剖面分布和季节动态与土壤可培养微生物数量一致。
除灌除草处理对土壤微生物数量和生物量的剖面分布特征产生了不同程度的影响。土壤细菌和真菌数量均以CK最大,RH次之,RS最小,而上壤放线菌数量则以RH最大,CK次之,RS最小；除灌除草处理对不同层次土壤微生物生物量的影响不同,其中,凋落物层土壤微生物生物量以RH最高,CK次之,RS最小,而0-5cm、5-10cm、10-15cm土层则以CK最高,RH次之,RS最低,除灌处理对土壤微生物生物量的影响大于除草处理,除草处理后土壤微生物生物量短期内有升高的趋势；除灌除草处理后,土壤微生物的剖面分布格局没有发生太大变化,但随土层加深,微生物数量和生物量递减的速率表现出差异,以RS最大,CK最小。
3.植物多样性对上壤酶活性的影响
不同林分类型的土壤蔗糖酶、脲酶、纤维素酶和脱氢酶活性表现出一定差异。云杉混交林各种酶活性均高于纯林；两类林分土壤酶活性均具明显表聚性,且有相似的季节动态,其中,蔗糖酶、脲酶与脱氢酶活性以秋季最低,夏季或春季最高,纤维素酶活性以夏季或秋季最高,春季最低。
除灌除草处理后,4种酶活性均降低,以CK>RH>RS,除灌处理的影响大于除草处理；4种酶活性均以CK随土层加深递减的速率最小,RS递减的速率最大；4种酶中,以纤维素酶活性随土层加深递减的速率最大,脱氢酶活性递减的速率最小。
4.植物多样性对凋落物分解的影响
不同林分类型的凋落物分解特征存在差异。云杉纯林凋落物分解速率低于混交林,约为混交林的65.51-99.63%,前期差异明显,后期差异较小。云杉纯林土壤微生物与小型土壤动物(主要是线虫和原生动物)对凋落物分解的贡献率达68.71-94.03%,大中型土壤动物对凋落物分解的贡献率为5.97～31.29%；云杉混交林土壤微生物与小型土壤动物对凋落物分解的贡献率达90.29～94.22%,大中型土壤动物对凋落物分解的贡献率为5.78-9.71%。各类土壤动物与微生物的共同作用大于仅微生物和小型湿生上壤动物的作用。在云杉凋落物分解的开始2年,除N元素出现富集外,P、K、Ca、Mg等元素均表现为释放,从释放量的大小来看,基本以混交林大于纯林。在该研究区域,所有分解者中以线虫、原生动物和真菌对凋落物分解起至关重要的作用。
除灌除草处理降低了凋落物分解速率,并且除灌处理导致的下降程度高于除草处理；各处理土壤动物与微生物对凋落物分解的共同作用大于仅微生物和小型湿生土壤动物的作用。各处理云杉凋落物分解的开始2年,N元素表现为富集,而P、K、Ca、Mg等元素则表现为释放。其中,除灌处理对凋落物养分释放表现为不利的影响,除Mg元素以外的N、P、K、Ca元素均以除灌处理释放量最小；除草处理在一定程度上有利于K元素的释放,而对N、P、Ca元素的释放影响不大,释放量仅次于或略等于对照。除灌除草处理后,凋落物分解过程中土壤动物、微生物及酶活性的变化并不是简单的以CK>RH>RS,而是表现出更加复杂的模式。由此认为,灌草层去除引起的凋落物组成和质量的改变,以及凋落物分解过程中土壤生物的多样性格局及酶活性的变化趋势是影响凋落物分解和养分周转的重要因素；同时,去除行为本身可能对森林生态系统的生物多样性以及物质循环功能产生干扰,干扰效应在处理早期比较明显,随时间推移逐渐减弱。
综上所述,植物多样性低的林分,其土壤生物多样性、土壤酶活性以及凋落物分解速率低于多样性高的林分；灌木层缺失对土壤生态过程的影响大于草本层缺失。因此,提高植物多样性,丰富凋落物组成,通过地上／地下生物互动,促进养分元素的循环,是增强生态系统各项功能的有效途径。
The relationship between biodiversity and ecosystem function is a hot and difficult problem in the field of ecological research, but a lot of studies are mainly concentrated on the research of grassland ecosystems, aquatic ecosystems and farmland ecosystem, because the means of research is inadequate, studies of forest ecosystem are relatively seldom, as compared with those ecosystems. Subalpine coniferous forests located in eastern edge of Qinghai-Tibet Plateau and the upper area of Yangtze River, it played an important and irreplaceable role in biodiversity conservation, water conservation and the global carbon cycle. Meanwhile, subalpine coniferous forests have some characteristics, for example, the hierarchical structure is clear and species richness is relatively simple, the law of update and succession is clearer; the type of structure and function is more various and so on. Those characteristics provide an ideal natural laboratory for the study of the composition of forest species and variations of forest ecosystems function and the process as affected by diversity change. A large number of studies have shown that the process is closely related to soil biodiversity, vegetation and ecosystems, interaction relationship of soil biological communities and vegetation determine ecosystem processes, the soil ecosystem is likely a bridge which makes biodiversity contact with ecosystem function. Above all, spruce artificial pure forest and mixed forest which were planted in1988were studied in this paper, the effects of plant diversity on soil microbes, soil enzyme activity, litter decomposition though field positioning observation and woodland means of control experiments and the influence of plant diversity on soil ecological process. Those results would enrich theoretical studies of the forest ecosystem biodiversity and ecosystem function, and provide basic data for restoration and reconstruction of subalpine degraded forest ecosystems. The main findings are as follows:
1. Effects of plant diversity on soil fauna
Different types of stand showed differences of the structure and spatial and temporal distribution of soil animal community. There are7197soil animals in Picea asperata pure plantation(not including protozoa, the same below); there are8831soil animals in Picea asperata mixed plantation. The number, density, biomass and diversity index of soil animal group of Picea asperata mixed plantation were higher than Picea asperata pure plantation. Among them, the number, density, biomass and diversity indices of macro-fauna have a significant or very significant difference. The quantities of saprophagous and predatory soil fauna in Picea asperata mixed plantation are higher than Picea asperata pure plantation, but the quantities of phytophagous soil animals are less than Picea asperata pure plantation. The ratio of A/C was effected by latitude and altitude, to some extent, reflect the characteristics of the distribution of soil animals in different zones; it is related to the heterogeneity of habitat, the degree of interference and diversity of plant communities in the same zone. The ratio of A/C of Picea asperata pure plantation was0.87, Picea asperata mixed plantation wasl.59and is about1.83times former. Soil protozoa density of Picea asperata pure plantation was365in1g fresh soil, soil protozoa density of spruce mixed forest was1046in1g fresh soil was significantly higher than Picea asperata pure plantation. The profile distribution of soil fauna of two stand types both have apparent surface gathering, and similar seasonal dynamics, are higher in summer than in spring and autumn.
Structure and distribution of soil animal communities showed different characteristics among different treatments.7529soil animals were collected in herb removal treatment (RH); the quantities of soil animals were6103in shrub removal treatment (RS).10986soil animals were collected in CK treatment. Through RH and RS treatment, the groups of soil macro-fauna showed increasing trend in short term, but the number of micro-and meso-soil fauna were decreasing; through RH and RS treatment, the density, biomass, and DG index of soil animals showed CK>RH>RS; number and proportion of saprophagous soil animals as well as A/C ratio were decreased, value of A/C was0.92in the RH treatment, value of A/C was0.82in the RS treatment, value of A/C of CK treatment was1.43; Through RH and RS treatment, soil protozoa density decreased,there were958soil animals in1g of fresh soil in RH treatment, there were1510soil animals in1g of fresh soil in RS treatment, there were2666soil animals in CK treatment, there was a significant difference among treatments; through RH and RS treatment, the profile distribution of soil animals has not changed, but a decreasing rate showed RS>RH>CK. Overall, effects of RS treatment on soil animal communities were more than RH treatment.
2. The effects of plant diversity on soil microbes
There were different component, number, biomass and temporal and spatial distribution of soil microbes in different plant types. Bacteria had the largest number in Picea asperata pure plantation and Picea asperata mixed plantation compared with other microbes, the order was Bacteria>actinomycetes>fungi, but the fungi ratio in mixed plantation was higher than that in pure plantation. The total number of microbes in Picea asperata pure plantation was significantly lower than that in Picea asperata mixed plantation, and number of actinomycetes and fungi in pure plantation were extremely lower than that in mixed plantation. Bacteria, fungi, total soil microbe number and actinomycetes in pure plantation in surface soil layer was higher than that in other soil layers, the actinomycetes had the lowest number in litter layer, and highest number in0-5cm layer compared with other soil layers. The order of Biomass in Picea asperata pure and mixed plantation was summer>spring>autumn. Biomass of soil microbes is the easy to decompose part in soil organic matter, which is an important source and sink, the profile distribution and seasonal dynamics of C/N of microbe biomass had the same trends with microbes number, and Picea asperata mixed plantation higher than that in Picea asperata pure plantation.
After shrub removal, soil microbes number, biomass and profile distribution changed a lot. The number order of soil bacteria and fungi was CK>RH>RS, and the number order of actinomycetes was RH>CK>RS. There was different affection on the microbes'biomass of soil layers after shrub and herb removal. The order of microbes'biomass in litter layer was RH>CK>RS, and The order of microbes'biomass in soil layers were CK>RH>RS, microbes' biomass had negative response after shrub removal, and had positive response after herb removal after a short period. There no huge changes in microbe profile distribution after shrub and herb removal, but microbe number and biomass reduced with the soil depth increased, the highest was RS, and the lowest was CK.
3. The effects of plant diversity on soil soil enzyme activity
There were diffrent activity of soil invertase, urease, cellulase and dehydrogenase in different plant types. All enzyme activity in the Picea asperata mixed plantation was higher than that in pure plantation. And enzyme activity in surface soil layer was higher than that in other soil layers, and had the same seasonal trends. The order of invertase, urease and dehydrogenase activity was autumn
After shrub cutting,4enzymes'activity reduced, the order was CK>RH>RS, and the enzymes'activity effected by removing shrub are higher than that of removing herb. The lowest degressive rate of4enzymes activity with the soil depth increased was CK, the highest was RS. And cellulase had the highest degressive rate with the soil depth increased among the4enzymes, and dehydrogenase had the lowest degressive rate.
4. The effects of plant diversity on litter decomposition
Litter decomposition had different characteristics in the different plantation. The decomposition rate of Picea asperata pure plantation was lower than that in the Picea asperata mixed plantation, decomposition ratio of Picea asperata pure plantation and mixed plantation is65.51～99.63%. the significant difference was found in early period, and there was tiny difference in later period. The contribution of soil microbes and meso-fauna (mainly nematode and protozoa) to litter decomposition in Picea asperata pure plantation are68.71～94.03%, contribution of soil macro-fauna to litter decomposition was5.97～31.29%; The contribution of soil microbes and meso-fauna to litter decomposition in Picea asperata mixed plantation are90.29～94.22%, contribution of soil macro and meso-fauna to litter decomposition was5.78～9.71%. The interaction decomposition of soil microbes and soil fauna were higher that effect of pure soil microbes or soil fauna. The front2years' decomposition, P、K、Ca、Mg element showed releasing, but the N showed enrichment. Releasing quantity of mixed plantation was higher that in pure plantation.
After shrub and herb removal, the litter decomposition rate decreased, and litter decomposition rate decreased caused by shrub removal was higher than that in herb removal. The interaction decomposition of soil microbes and soil fauna were higher that effect of pure soil microbes or soil fauna. The front2years' decomposition, P、K、Ca、Mg element showed releasing, but the N showed enrichment. Shrub removal had negative effect on the litter decomposition, N、P、K、Ca element showed releasing excepting Mg. Herb removal had positive effect on K releasing, and had little affection on the N、P、Ca releasing, the releasing quantity had no significant difference compared with CK. After shrub and herb removal, the changes of soil fauna, soil microbes and enzyme activity during the litter decomposition hadn't the order CK>RH>RS, but had more complicated models. The result can be conclude that the shrub and herb removal lead to the composition and quality changes, and the soil fauna diversity and the change trends of enzyme were the main factors which lead to litter decomposition and nutrition cycle during the litter decomposition. Meanwhile, the removal is an important interference which affects the biodiversity and material cycle; the interference affection was significant in early period.
To summary, compared with rich plant diversity plantation, the soil biodiversity, soil enzyme activity, and litter decomposition rate were lower than that in the poor plant diversity plantation. Stronger affection on the soil ecology process was found in shrub removal compared with herb removal. Therefore, to increase plant diversity, to enrich litter composition, to interact upper and underground organisms, to improve nutrition recycles, can enhanced the functions of forest ecosystem.
引文
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