长白落叶松（Larix olgensis）工业人工林的模拟间伐

应用马尔柯夫过程理论,在获得林分直径转移概率的基础上,采用间伐最小径阶林木,最大径阶林木,中间径阶林木三种间伐方式,在保留不同密度情况下对长白落叶松工业人工林进行模拟间伐,提出了适宜的保留密度和相应的抚育间伐对象。结果表明：马尔柯夫过程确能反映长白落叶松工业人工林的直径转移过程,利用马尔可夫过程理论对长白落叶松工业人工林进行模拟间伐实现了依据培育时间来确定间伐方法和措施,提高了长白落叶松工业人工林经营管理的精准性;长白落叶松工业人工林成林后的间伐无论从培育森林方面,还是从取得木材、加大林分收益方面考虑,都应该以间伐小径阶的林木为主,注重培育I、II级木,间伐III, IV级木;20～25 a长白落叶松工业人工林间伐后的保留经营密度以0.7为宜。     

天山雪岭云杉林粗木质残体储量特征

粗木质残体(coarse woody debris,CWD)在天山雪岭云杉林生态系统中起着重要的结构性和生物地球化学作用,解释其储量特征是研究CWD的基础,但尚未有大尺度研究见诸报道。以天山雪岭云杉8 hm~2森林动态监测样地为研究对象,采用野外调查、室内试验以及数据分析相结合的方法,调查了样地内CWD的储量组成、径级以及分解等级分布格局等基本特征及其影响因子。结果表明:(1)天山雪岭云杉8 hm~2森林动态监测样地内共有直径≥10 cm的CWD 936株,CWD的密度、体积、储量分别为117株/hm~2,15.13 m~3/hm~2,4.52 t/hm~2;其中倒木是CWD的主要贡献者,占CWD总储量的52.21%;(2)样地内各径级CWD的数量呈典型的倒"J"型结构,直径30 cm的CWD个体占全部CWD的83%;(3)样地内CWD总体上处于以Ⅱ、Ⅲ分解等级为主的中度分解状态,CWD径级越大,分解程度越高;(4)林分密度、郁闭度和海拔是影响天山雪岭云杉林CWD储量特征的主要因素。研究可为天山雪岭云杉林的可持续发展与经营提供科学依据。     

樟子松人工林一级枝条基径和枝长模型的研究

以东北林业大学帽儿山实验林场樟子松人工林为研究对象,采用枝解析的方法,于2002年、2003年测定了53株林木（年龄17~38 a,直径8.61~21.5 cm,树高7.48~18.24 m）的树冠变量,建立了基于总着枝深度（DINC）和树冠内一级枝条基径（BD）、枝长（BL）的预估模型。对于大小相同树木的一级枝条, 这些树冠变量随着DINC的增加而增大,而林木的胸径（DBH）、树高（HT）变量又很好地反映了不同大小树木的基径和枝长的变化。采用独立检验样本对构建的树冠内一级枝条基径和枝长模型进行了拟合统计量和精度检验,结果表明：模型预测效果良好,精度均达到95%以上。构建的一级枝条基径和枝长模型为进一步合理地描述樟子松人工林树冠的形状及其变化以及三维可视化经营提供依据。     

樟子松沙地人工林直径分布模拟

利用反 Weibull函数模拟了章古台沙地樟子松人工林的直径分布规律 .结果表明 ,反 Weibull分布函数模拟樟子松人工林的直径分布效果好 ,具有精度高、适应性强等特点 .分别建立了反 Weibull分布函数 3个参数与林分年龄及密度之间的关系方程以及直径分布与 3个参数的回归方程 ,这些方程可用来预估林分产量、出材量以及评价经营效果     

樟子松人工林一级枝条基径和枝长模型的研究

以东北林业大学帽儿山实验林场樟子松人工林为研究对象,采用枝解析的方法,于2002年、2003年测定了53株林木（年龄17～38a,直径8．61～21．5cm,树高7．48～18．24m）的树冠变量,建立了基于总着枝深度（DINC）和树冠内一级枝条基径（BD）、枝长（BL）的预估模型。对于大小相同树木的一级枝条,这些树冠变量随着DINC的增加而增大,而林木的胸径（DBH）、树高（HT）变量又很好地反映了不同大小树木的基径和枝长的变化。采用独立检验样本对构建的树冠内一级枝条基径和枝长模型进行了拟合统计量和精度检验,结果表明：模型预测效果良好,精度均达到95％以上。构建的一级枝条基径和枝长模型为进一步合理地描述樟子松人工林树冠的形状及其变化以及三维可视化经营提供依据。     

不同密度樟子松人工林土壤碳氮磷化学计量特征

以科尔沁沙地不同密度(490、750、1550、1930、2560株·hm^-2)樟子松人工林(栽植于1980年)为研究对象,分析林分密度对土壤碳、氮、磷浓度及其计量比的影响,研究林分密度与土壤养分状况的关系。结果表明:随着樟子松林密度增加,各土层(0~10、10~20和20~40 cm)土壤有机碳、全氮、全磷浓度和C∶N呈先增加后降低趋势,而土壤有效磷浓度呈先降低后增加趋势。土壤有机碳浓度在490株·hm^-2密度小于其他密度,而有效磷浓度大于其他密度;土壤C∶P和N∶P在2560株·hm^-2密度显著大于其他密度。各密度樟子松林土壤有机碳、全氮、全磷和有效磷浓度在0~10 cm土层显著大于10~20和20~40cm土层,樟子松人工林土壤养分具有表聚性。通过典范对应分析发现,密度对樟子松林土壤养分影响的主要因子是土壤有机碳、全氮和全磷,且密度为1550株·hm^-2时土壤有机碳、全氮、全磷和碱解氮浓度较高,而C∶P和N∶P较低。因此,当樟子松人工林密度为1550株·hm^-2时,土壤养分浓度较高,林木生长较好,为最佳经营密度。     

不同密度樟子松人工林土壤碳氮磷化学计量特征

以科尔沁沙地不同密度(490、750、1550、1930、2560株·hm-2)樟子松人工林(栽植于1980年)为研究对象,分析林分密度对土壤碳、氮、磷浓度及其计量比的影响,研究林分密度与土壤养分状况的关系。结果表明:随着樟子松林密度增加,各土层(0～10、10～20和20～40 cm)土壤有机碳、全氮、全磷浓度和C∶N呈先增加后降低趋势,而土壤有效磷浓度呈先降低后增加趋势。土壤有机碳浓度在490株·hm-2密度小于其他密度,而有效磷浓度大于其他密度;土壤C∶P和N∶P在2560株·hm-2密度显著大于其他密度。各密度樟子松林土壤有机碳、全氮、全磷和有效磷浓度在0～10 cm土层显著大于10～20和20～40cm土层,樟子松人工林土壤养分具有表聚性。通过典范对应分析发现,密度对樟子松林土壤养分影响的主要因子是土壤有机碳、全氮和全磷,且密度为1550株·hm-2时土壤有机碳、全氮、全磷和碱解氮浓度较高,而C∶P和N∶P较低。因此,当樟子松人工林密度为1550株·hm-2时,土壤养分浓度较高,林木生长较好,为最佳经营密度。     

江西武夷山中亚热带南方铁杉针阔混交林群落组成与结构

长期监测型固定样地是研究物种分布格局、群落动态和生物多样性维持机制等森林生态系统特征、过程与机理的重要平台。我国亚热带地区已建立的森林固定样地多为阔叶林类型,而针叶(阔叶混交)林类型十分有限。按照巴拿马Barro Colorado Island大型森林固定样地的建设规范,于2014年在江西武夷山国家级自然保护区内海拔1800 m左右的南方铁杉(Tsuga chinensis var. tchekiangensis)天然种群分布区域建立中亚热带针阔混交林6.4 hm~2固定样地。木本植物调查结果表明:(1)样地内胸径≥1 cm的木本植物共有29科53属89种,显著低于我国亚热带阔叶林样地平均水平,但显著高于温带针叶(阔叶混交)林样地平均水平;(2)在区系组成上,科以热带成分为主,而属以温带成分为主;(3)样地内独立个体数密度为2252株/hm~2,与温带针叶(阔叶混交)林平均水平相当,但显著低于亚热带阔叶林平均水平;(4)胸高断面积为37.89 m~2/hm~2,与亚热带阔叶林样地平均水平相当,但显著低于温带针叶(阔叶混交)林平均水平;(5)群落成层现象显著,优势种明显,多度排名前4位的物种的个体数占总个体数的55%,而排名后40的物种仅占1%;(6)群落总径级结构呈倒"J"型分布,其中胸径≤10 cm的小径木占总个体数的76.9%,而胸径30 cm的大径木仅占5.3%;(7)主要树种的径级结构有偏正态分布和"L"型分布等类型,但它们的种群在空间上均呈现显著的聚集分布特征;(8)南方铁杉虽然是群落现阶段最重要的优势种,但它的种群更新缓慢。研究结果充分显示该样地对中国森林多样性监测以及南方铁杉种群保育的重要意义。     

大花序桉心边材变异规律和候选基因挖掘研究

为探究大花序桉(Eucalyptus cloeziana)心材比例差异显著的不同家系间心边材变异规律,挖掘心材变异相关的候选基因,为珍贵用材树种高效培育及育种利用提供基因资源。以18 a生的2个心材比例差异显著的大花序桉家系为材料(家系1和2),各制作解析木3株,沿着树干以1 m为区间分段截取圆盘,测量东西和南北2个方向的带皮直径、去皮直径、总年轮数、边材年轮数、边材直径,并开展心材和边材径向和轴向分析。同时利用各解析木胸径处初生木质部样品进行DNA混池测序,发掘等位基因频率差异显著的SNP位点并挖掘相关功能基因。结果表明,大花序桉边材宽度和心材半径的方位变异中家系2大于家系1,平均差值分别为0.7和5.5 cm,在随树高的变异中,家系1和2的心材半径和心材年轮数的下降速率分别为0.40和0.64及0.43和0.36。两家系间基本密度差异显著,家系1为0.80～0.82 g/cm³,家系2为0.75～0.78 g/cm³。基本密度与树高、横截面半径和心材半径呈显著负相关,与顺纹抗拉强度、弦面硬度和部分力学性质呈显著正相关。利用DNA混池测序共...     

棕背(鼠平)、红背(鼠平)栖息地选择的研究

棕背(鼠平)(Clethrionomys rufocanus)与红背(鼠平)(Clethrionomys rutitus)是吉林省林木主要害鼠,本文应用数量化方法,分析野外调查数据,阐明它们对栖息地选择的生物学特性,为森林鼠害防治工作提供依据。 1.研究方法 1983年9月15日至10月15日,吉林省林业厅在全省43个县同时对森林害鼠的种类、分布密度和危害进行调查,共布41376铗日,捕鼠2271只。本文选择东部山区樟子松人工林31块、红松人工林50块、落叶松人工林78块、人工针叶混交林15块、天然次生林42块样地,对棕背(鼠平)与红背(鼠平)的铗日捕获     

樟子松人工林树冠表面积及体积预估模型的研究

基于樟子松(Pinus sylvestris var. mongolica)人工林6块固定标准地30株枝解析数据,在分析树冠表面积和树冠体积与林分变量和林木变量的基础上,利用幂函数建立了树冠表面积（CSA）和树冠体积(CV)的预估模型,同时还对林木材积生长量与CSA和CV进行了相关分析。研究结果表明：樟子松人工林树冠表面积和树冠体积随着林木胸径、树高和冠长的增大而增大,林木材积生长量与树冠表面积和树冠体积均明显呈线性关系。不同林分条件的樟子松人工林CSA和CV随林分年龄和胸径的增大而增大,CSA随林分密度的增大而减小,而CV与林分密度相关不紧密。林分树冠表面积和树冠体积预估模型的检验结果表明,两个模型的平均相对误差都在±8%之内,预估精度均大于91%,说明所建模型可以很好地预估樟子松人工林不同林分条件下的林木树冠表面积和树冠体积。     

Tree rings as criteria for selection of timber for building of chapels in the Republic of Karelia

Data on the quality of timber used for building chapels – small buildings for public worship, enabled determination of visual criteria used for the selection of timber in the Republic of Karelia, located in the north-western part of the Russian Federation.It was revealed that as for the majority of timber structures in the region, Scots pine (Pinus sylvestris L.) logs were selected for the chapels. Trees perceived as 'Holy' such as spruce (Picea abies Kr.), were not used in these worship structures, in contrast to practices in the neighboring Leningrad Region. The age of the logged trees was 100–150 years, with a high proportion of young trees.Data obtained proved convincingly that compared with huge parish churches, selection of timber for these small chapels was not so strict. Multivariate analysis of variance showed that an ethnological factor played a significant role in the selection of wood for the buildings. Mature trees with narrow tree rings were preferred for worship structures preserved in villages populated by Karelians – the Baltic-Finnic tribe living in the region since ancient times.In villages with a dominantly Russian population, young pines with wide annual rings were dominantly used. A correlation between the height of the structures and the average diameter of the logs used was found with the coefficient of determination high in Karelian villages and lower in Russian villages.     

A Functional and Structural Mongolian Scots Pine (Pinus sylvestris var. mongolica) Model Integrating Architecture, Biomass and Effects of Precipitation

Mongolian Scots pine (Pinus sylvestris var. mongolica) is one of the principal tree species in the network of Three-North Shelterbelt for windbreak and sand stabilisation in China. The functions of shelterbelts are highly correlated with the architecture and eco-physiological processes of individual tree. Thus, model-assisted analysis of canopy architecture and function dynamic in Mongolian Scots pine is of value for better understanding its role and behaviour within shelterbelt ecosystems in these arid and semiarid regions. We present here a single-tree functional and structural model, derived from the GreenLab model, which is adapted for young Mongolian Scots pines by incorporation of plant biomass production, allocation, allometric rules and soil water dynamics. The model is calibrated and validated based on experimental measurements taken on Mongolian Scots pines in 2007 and 2006 under local meteorological conditions. Measurements include plant biomass, topology and geometry, as well as soil attributes and standard meteorological data. After calibration, the model allows reconstruction of three-dimensional (3D) canopy architecture and biomass dynamics for trees from one- to six-year-old at the same site using meteorological data for the six years from 2001 to 2006. Sensitivity analysis indicates that rainfall variation has more influence on biomass increment than on architecture, and the internode and needle compartments and the aboveground biomass respond linearly to increases in precipitation. Sensitivity analysis also shows that the balance between internode and needle growth varies only slightly within the range of precipitations considered here. The model is expected to be used to investigate the growth of Mongolian Scots pines in other regions with different soils and climates.     

三种温带森林大型土壤动物群落结构的时空动态

对帽儿山3种典型森林群落大型土壤动物进行了连续6个月的野外调查研究。通过系统分析,共获得大型土壤动物3604只,隶属于3门6纲17目50科。其中正蚓科(Lumbricidae)、线蚓科(Enchytraeidae)和石蜈蚣目(Lithobiomorpha)为优势类群,常见类群11类。结果表明:(1)水平分布上,密度和生物量红松人工林最高,其次为硬阔叶林,蒙古栎林最少;类群数硬阔叶林最多,蒙古栎林最少;香农指数和丰富度指数均为蒙古栎林最高,红松人工林最低;优势度指数与两者相反;均匀度指数蒙古栎林最高,硬阔叶林最低;(2)垂直分布上,个体密度、类群数及生物量均差异显著(P < 0.001)。3个样地大型土壤动物个体密度表聚性明显;类群数红松人工林自凋落物层向下减少,硬阔叶林和蒙古栎林0-10 cm最多;生物量在0-10 cm土层最大;香农指数随深度增加而减小,优势度指数则相反;(3)在时间变化上,5月和10月个体密度和类群数较多,9月生物量最大;香农指数和优势度指数差异显著(P < 0.01),其他指数各月间无明显差异;(4)与土壤环境因子关系上,总有机碳含量与类群数、个体密度及生物量显著正相关,容重与香农指数显著负相关;典型对应分析结果表明,不同类群大型土壤动物与环境相关性不同。     

长白山阔叶红松林退化生态系统的土壤呼吸作用

选择处于全球变化中国东北样带东部典型生态系统的长白山阔叶红松林作为研究区,采用动态气室-CO₂红外分析法测定了森林生态系统不同退化阶段的土壤呼吸作用.结果表明：在生长季,长白山阔叶红松林不同退化阶段的土壤呼吸动态变化呈单峰型曲线,在7-8月达到最大值;不同退化阶段林地土壤呼吸大小顺序为：杨桦林＞蒙古栎林＞阔叶红松林＞硬阔叶林＞裸地.其中,杨桦林和蒙古栎林样地的碳释放量分别为对照阔叶红松林的1.4和1.3倍,硬阔叶林和裸地的碳释放量分别为对照阔叶红松林的88%和78%.     

The contribution of stored water to transpiration in Scots pine

Abstract. The amount of water available diurnally and annually from the storage tissues was measured in plots of Scots pine trees with four different population densities (608–3281 trees per ha) in a 40-year-old plantation in north eastern Scotland. The water storage capacity of stems, branches, and foliage was estimated from equations derived from harvested trees and measurements of relative water content. On average 64% of the water considered to be available for transpiration was in the stem sapwood and less than 5% in the phloem, cambium and foliage. Trees on the plot with the highest population density had a water storage capacity of 212 m3 ha^?1 (21.2 mm), whereas those on the plot with the lowest population density had a water storage capacity of 124 m3 ha^?1 (12.4 mm). The utilization of stored water in transpiration was estimated from seasonal and diurnal measurements of the relative water content of foliage and stem sapwood. The largest change in sapwood relative water content over a 2-week period was a reduction of 27% corresponding to extraction from the sapwood of 2.5 and 5.1 mm of water on the plots with the lowest and highest population densities, respectively. In rapidly changing weather conditions 1–1.5 mm day^?1 could be removed from the stem sapwood alone. Since transpiration rarely exceeded 3 mm day^?1, 30–50% of the transpired water was extracted from water stored in the stem sapwood over short periods. Trees on the plot with the lowest population density occasionally had slightly higher relative water contents and exhibited larger diurnal fluctuations than those on the plot with the highest population density, possibly because of differences in wood density. Sapwood water content was generally lower at times of high transpiration rate and in winter during freezing conditions. Resaturation took several months to complete during the winter.     

Lack of regeneration and climatic vulnerability to fire of Scots pine may induce vegetation shifts at the southern edge of its distribution

Aim Forest ecosystems dominated by fire‐sensitive species could suffer shifts in composition under altered crown fire regimes mediated by climate change. The aims of this study were to: (1) study the spatio‐temporal patterns and the climatic distribution of fires in Scots pine (Pinus sylvestris) forests during the last 31 years in north‐eastern Spain, (2) evaluate the climatic vulnerability to fire of these forests in Spain, (3) analyse the regeneration of Scots pine after fire, and (4) predict the mid‐term maintenance or replacement of Scots pine in burned areas. Location Catalonia (north‐eastern Spain): the southern distribution limit of Scots pine. Methods We characterized the spatio‐temporal and the climatic distribution of fires that occurred in Catalonia between 1979 and 2009. We used a generalized linear model to characterize the climatic vulnerability to fire of Scots pine in the whole of Spain. We assessed the regeneration of the species after crown fires in nine burned areas in Catalonia. The resulting data were integrated into a stochastic matrix model to predict the mid‐term maintenance or replacement of Scots pine in burned areas. Results During the last three decades, Scots pine forests distributed in dry sites were most affected by fire. Our assessment of the vulnerability to fire of Scots pine forests in Spain as a whole, based on climatic and topographical variables, showed that 32% of these forests are vulnerable to fire, and that this proportion could increase to 66% under a conservative climate change scenario. Field data showed almost no regeneration of Scots pine after crown fires, and a limited capacity to recolonize from unburned edges, even in relatively old fires, with 90% of recruits located in the first 25 m from the edge. This process could be delayed by the elapsed time for new recruits to achieve reproductive maturity, which we estimated to be c. 15 years. Finally, our matrix model predicted the replacement of burned Scots pine forests by oak (Quercus sp.) forests, shrublands or mixed resprouter forests. Main conclusions Increased vulnerability to fire of Scots pine forests under future, warmer conditions may result in vegetation shifts at the southern edge of the distribution of the species.     

Genetic structure of montane isolates of Pinus sylvestris L. in a Mediterranean refugial area

Aim This work investigates the population genetic effects of periodic altitudinal migrations and interstadial fragmentation episodes in long‐term Scots pine (Pinus sylvestris L.) populations at a regional scale. Location The study focuses on Scots pine populations in the northern Meseta and peripheral mountain chains, central and north‐western Iberian Peninsula. The ample macrofossil record in the area shows that this 60,000‐km² region represent a glacial refugium for Scots pine. The species occupied large areas on the Meseta plains during glacial cold stages, but it has periodically sheltered at high elevation in the surrounding mountain chains during warm episodes, conforming to a fragmented pattern similar to its present‐day distribution. Methods We perform a fine‐scale chloroplast microsatellite (cpSSR) survey to assess the genetic structure of 13 montane Scots pine isolates in the northern Meseta (total N = 322 individuals). Using a hierarchical analysis of molecular variance (amova ), we test the hypothesis of genetic isolation among disjunct mountain areas. We use a standard coalescence model to estimate genealogical relationship among populations, investigating the potential role of the regional relief as a factor influencing historic gene exchange among Scots pine populations. Results Population haplotypic diversity was high among Scots pine populations (H_e = 0.978), greater than values reported for other more thermophilic pine species in the Iberian Peninsula. The amova revealed low (but significant) differentiation among populations (Φ_ST = 0.031, P = 0.010), showed that the disjoint montane distribution could not account for the genetic divergence among areas (Φ_CT = 0.012, P = 0.253), and that there was non‐trivial subdivision among populations within the same mountain region (Φ_SC = 0.021, P = 0.012). The genealogical relationships among populations showed that Scots pine isolates growing on disjoint mountain blocks, but on slopes flowing to the same basin, were genetically closer than populations growing on different slopes of the same mountain chain, flowing to different basins. Main conclusions The observed genetic structure for Scots pine is consistent with its population history, inferred from the palaeobotanical record, with vertical migrations throughout climatic pulses and with the drainage basins and large long‐term population sizes connecting different mountain blocks during the cooler glacial periods. Overall, the results suggest that, despite periodic interstadial fragmentation episodes, Scots pine biology provides for the long‐term maintenance of high within‐population and low among‐population genetic diversity at neutral genetic markers.     

Population and reproductive structure of an endangered juniper from coastal pine forests

Plant species adapted to coastal systems are suffering severe declines. This is the case of the critically endangered Juniperus navicularis, which colonizes fossil dunes in the southwestern Iberian Peninsula and grows in the understory of pine plantations. This endemic juniper presents low reproduction rates and a decreasing number of populations has been reported in its western distribution edge. However, age structure and the demographic situation of its easternmost populations, that have been recently discovered, remain unknown. We used dendrochronological procedures to estimate the age structure of the easternmost isolated population of J. navicularis, considering three subpopulations under contrasting management regimes and selecting plots under pine cover and in open areas. We also assessed sex ratio, age at maturity and reproductive effort. We observed the predominance of 20–40 years-old stems in all study plots and a remarkable lack of young stems in open areas. Stem density was higher under pine canopies. There, junipers were, on average, younger than pine trees, meaning that junipers had grown in already forested areas. Only one sex per plot was found in all study plots but one, in which no reproductive stems were recorded, suggesting the existence of clonal growth. Reproductive effort increased with age in females, but was independent in males. We hypothesize that clonal growth might occur as a survival strategy to endure the harsher conditions of distribution edges. These results will help to develop more effective management strategies to preserve this endangered species, contributing to our understanding of dynamics of coastal ecosystems and setting the basis for future genetic studies.     

帽儿山3种森林生态系统土壤动物与土壤呼吸及其相互关系研究

土壤呼吸是森林生态系统碳循环的关键过程,土壤动物可通过自身代谢及影响微生物活动调控土壤呼吸,因此研究土壤动物与土壤呼吸的相互关系对进一步揭示生态系统碳循环的规律和机理具有重要意义。通过野外定点,以帽儿山3种森林生态系统的土壤呼吸及土壤动物为研究对象,探讨不同森林生态系统的土壤呼吸、土壤动物个体密度和生物量的时间变化规律及二者相互关系。结果表明:(1)3种森林生态系统土壤总呼吸速率与土壤异养呼吸速率均呈现先增强后减弱的时间动态变化(P<0.05),且不同森林生态系统土壤异养呼吸速率差异显著(P<0.05),表现为硬阔叶林最高,红松人工林最低;(2)3种森林生态系统土壤动物生物量也具有显著的时间动态变化(P<0.05),均在9月份达到最大,且不同森林生态系统土壤动物个体密度显著不同(P<0.05),蒙古栎林土壤动物个体密度显著小于红松人工林与硬阔叶林;(3)通过回归分析可得,土壤动物数量及生物量的增加抑制了土壤呼吸速率,尤其在生长季初期、末期。研究表明土壤动物可通过抑制微生物生命活动和降低根系呼吸从而对土壤总呼吸及异养呼吸产生负反馈作用,三者是不可分割的整体,与土壤温度、水分等环境因子共同调控着土壤呼吸。