生长空间竞争指数及其在油松、侧柏种内竞争中的应用研究

研究林木个体之间的竞争是研究森林生态系统基础,林木竞争的研究方法主要有定性描述和定量分析两种方法并先后提出了20多种衡量竞争程度的竞争指数系统,但有关学者研究后认为,这些竞争指数都没有较好地解决了定量分析中树冠变化的问题。论文首次提出生长空间竞争指数(Growthspacecompetitionindex,GSCI),系树冠表面积与胸高断面积之比,由于树冠外围面积代表树冠接受光照最强的部分并且是决定生产效率的重要指标,因此在群体和群落的竞争中作为反映竞争参数,较好地解决了定量分析中树冠变化的问题,为树冠覆盖程度的无量纲度量。本文还在油松(Pinus tabulaeformis)、侧柏(Platycladus orientalis)林种内个体竞争中得到应用,与常用的简单竞争指数(Competition index,CI)和空间竞争指数(Growth space index,GSI)进行了对比研究。     

油松、侧柏林种内竞争特点的对比研究

植物竞争研究一直是种群生态学最活跃的研究领域之一,论文以林木个体相对胸径、相对树高、相对冠幅、相对冠长、相对冠面积、树冠体积、树冠伸展度、冠长率、圆满度、投影比、生长空间指标、生长空间竞争指数和简单竞争指数13个变量作为对比研究油松、侧柏林个体之间竞争关系的指数,采用这些指标的变异系数对其种内竞争的差异程度进行比较研究,并用这些变量作为评价指标将其竞争能力分为两类:种内竞争性强的树种和种内竞争性弱的树种。油松为种内竞争性强的树种,而侧柏为种内竞争性弱的树种,已有的研究结果已从生物量、根系特征、抗旱机理、耗水特性、分布及结构特点五个方面为这种划分提供了理论支持。     

Effect of climate and competition on radial growth of Pinus massoniana and Schima superba in China’s subtropical monsoon mixed forest

Tree growth is affected by many exogenous and endogenous factors, especially climate and competition. To address the issue of how these factors influence tree growth under global warming, dendroclimatology combined with competition analysis was used to examine the radial growth response of two major species – Pinus massoniana and Schima superba in subtropical monsoon mixed forests in southern China. For growth-climate relationship analysis, residual and ARSTAN chronologies of these two species were compared for the response function analysis over the past several decades. In terms of competition analysis, annual basal area increment (BAI) over several time intervals was calculated and linear mixed model techniques were employed. Several distance-independent competition indices were measured, including density (N), basal area (BA), and sum of diameter at breast height (SDBH). For competition trees with greater diameter (GR) than the subject tree, NGR, BAGR, SDBHGR were also calculated. Growth-climate relationship analysis suggested that tree growth varied in response to climate and was mostly correlated with minimum, mean and maximum air temperature as indicated by the adjusted R² value. Summer and winter temperatures had a negative effect on tree growth, while early spring temperature showed positive effects. Competition analysis suggested that subtropical trees had been affected by competition in varying degrees. Trees in the oldest forest had the most competition stress, while trees in younger forests were less affected by competition according to the adjusted R² value. Given that subtropical forests in China have been found to have more smaller trees with recent warming, it is expected that these forests will experience a decreasing effect of competition in near future. This can have profound effects on forest productivity and carbon sequestration potentials in Chinese subtropical forests.     

Importance and intensity of competition along a fertility gradient and across species

Questions: 1. Can the importance and the intensity of competition vary independently along a nutrient gradient? 2. Are these variations species dependent? Location: Sub‐alpine pastures of the northern French Alps. Methods: Competition intensity measures how much competition decreases the performances of an organism. Competition importance measures how much competition contributes to affect performance, among other processes (such as environmental stress or disturbance). Competition intensity and importance were measured on three co‐occurring species: Festuca rubra, a perennial grass, and two forbs of contrasting basal area, Chaerophyllum hirsutum and Alchemilla xantho‐chlora. A neighbour removal experiment was performed on Festuca rubra in three sub‐alpine grassland communities differing in fertility and on Chaerophyllum hirsutum and Alchemilla xanthochlora in the two more fertile of these communities. The importance of competition was quantified using an index proposed by Brooker et al. (2005). Results: Competition intensity and importance showed different patterns of variation along the fertility gradient for Festuca rubra: competition importance decreased with decreasing fertility whereas competition intensity did not change. The largest forb was the least affected by competition. Our results suggest that the importance of competition for all three species depended on their individual tolerance to low nutrient availability. Conclusions: 1. The distinction between the importance and the intensity of competition is helpful to explain conflicting results obtained on the variations of competition indices along productivity gradients. 2. The choice of a phytometer can affect the conclusions drawn from empirical studies.     

临安区不同森林类型竞争指数比较研究

竞争对树木生长有重要影响,通常用竞争指数定量描述树木之间的竞争关系。以浙江省杭州市临安区的乔木林为对象,利用2004年临安区森林资源监测数据,基于Voronoi图的Hegyi竞争指数,在区域尺度上对临安区多种森林类型的竞争关系进行分析。结果表明：在单木水平上,不同森林类型单木的总竞争指数、种内竞争指数、种间竞争指数在胸径5-10 cm最大;单木平均竞争指数随着胸径的增大而降低,胸径 > 30 cm的竞争压力得以缓解;临安区林木的竞争主要来源于种内竞争,天然林的种内竞争占总竞争指数的比例 > 50%,人工林的种内竞争占总竞争指数的比例 > 60%。径级平均总竞争指数、种内竞争指数、种间竞争指数与胸径的关系均服从对数函数关系。在林分水平上,不同森林类型的林分总竞争指数、种内竞争指数、种间竞争指数存在显著差异。平均总竞争顺序为：天然阔叶林 > 天然针阔混交林 > 人工针叶林 > 天然针叶林 > 人工针阔混交林 > 人工阔叶林。种内平均竞争顺序为：人工针叶林 > 天然阔叶林 > 天然针叶林 > 天然针阔混交林 > 人工针阔混交林 > 人工阔叶林。种间平均竞争顺序为：天然针阔混交林 > 人工针阔混交林 > 天然阔叶林 > 天然针叶林 > 人工针叶林 > 人工阔叶林。人工阔叶林的平均总竞争指数、种内竞争指数和种间竞争指数均为最小,该森林类型以山核桃经济树种为优势树种,说明为提高山核桃产量,人为控制低种植密度的经营措施有效降低了林分竞争强度。     

西双版纳热带季节雨林林冠种群种内、种间竞争的初步研究

利用Hegyi的竞争指数模型,分析了西双版纳热带季节雨林的主要类型绒毛番龙眼群落中林冠种群种内和种间的竞争强度及种群和个体在群落中的竞争干扰状态.结果表明,季节雨林中林冠种群的种间竞争大于种内竞争,每个林冠种群一般与多个其它林冠种群发生竞争.多数种群既被竞争种干扰,又干扰竞争种.个体在群落中所处的层次越高,受到的竞争干扰越小,下层中的个体多数处于受压迫状态.群落中没有完全的竞争优胜者.只有成年个体处于竞争优势地位,幼苗、幼树又能在受压迫中生存的种群,才能在群落中保持长期存在.     

Effects of individual size,local competition and canopy closure on the stem volume growth in a monoclonal Japanese cedar (<Emphasis Type="Italic">Cryptomeria japonica</Emphasis> D. Don) plantation

We studied how the dominant factor affecting stem volume growth changes during stand development in a monoclonal stand of Cryptomeria japonica D. Don. Stem analysis was used to compare growth history of trees in an unthinned plot (closed canopy) and a thinned plot (open canopy). In the unthinned plot, the dominant factor affecting stem volume growth was basal area (BA) before canopy closure, whereas neighborhood competition index (CI) was the dominant factor after canopy closure. In contrast, the dominant factor affecting stem volume in the thinned plot was BA throughout stand development. Spearman’s rank correlation coefficient between BA and CI continued to increase after canopy closure and size rank among individuals became increasingly fixed. Our results indicated that stem volume growth shifts from size-dependent to competition-dependent growth at canopy closure. The apparent correlation between tree size and growth rate observed in many previous studies may be the result of competition-mediated positive feedback between size and growth.     

Growth models of silver birch (Betula pendula Roth.) on two volcanic mountains in the French Massif Central

Silver birch woodlands of two volcanic mountains (altitude 850 m and 1450 m) were studied in order to establish a growth model of birch. Height, radial increment and crown width were measured on both sites taking into account two situations: isolated birch or birch within a stand. For the latter case two categories were sampled considering the competition status of the tree: birch suffering the most severe competition (suppressed tree) were distinguished from trees facing the lowest competition (dominant tree). Measures of competition were also made using plots where each tree was located by its coordinates and its height, trunk circumference and crown width recorded. Examination of radial growth curves showed similar patterns for the two sites for the same category of tree. Radial growth was always inferior for the site located at the highest altitude but tree lifespan was about two-fold longer. Radial increment data were used to calculate circumference as function of tree age. Models predicting circumference with time were then established for each category of tree (dominant, suppressed or isolated) using Chapman–Richards'model. Height and crown width models were produced using circumference as the predictor. Competition indices based on vertical or horizontal angles weighted by the distance were calculated for birches in stands. Indices using vertical angles appeared to be more resistant and robust to characterize local competition. Competition index was then associated with the parameters of Chapman-Richards' growth circumference model for the two sites and models predicting the circumference from the age of the tree and its competition index are proposed.     

Plant competition for light analyzed with a multispecies canopy model

Summary A multispecies canopy photosynthesis simulation model was used to examine the importance of canopy structure in influencing light interception and carbon gain in mixed and pure stands of wheat (Triticum aestivum L.) and wild oat (Avena fatua L.), a common weedy competitor of wheat. In the mixtures, the fraction of the simulated canopy photosynthesis contributed by wheat was found to decline during the growing season and this decline was closely related to reductions in the amount of leaf area in upper canopy layers. For both species in mixture and in monoculture, simulated photosynthesis was greatest in the middle or upper-middle canopy layers and sensitivity analyses revealed that canopy photosynthesis was most sensitive to changes in leaf area and leaf inclination in these layers. Changes in LAI and leaf inclination affected canopy carbon gain differently for mixtures and monocultures, but the responses were not the same for the two species. Results from simulations where the structural characteristics of the two species were substituted indicated that species differences in leaf inclination, sheath area and the fraction of leaf area alive were of minor consequence compared with the differences in total leaf area in influencing relative canopy carbon gain in mixtures. Competition for light in these species mixtures appears to be influenced most by differences in the positioning of leaf area in upper canopy layers which determines, to a great extent, the amount of light intercepted.     

基于树冠重叠面积的天然混交林林木竞争指数

构建基于树冠重叠面积和林木混交关系的竞争指数,探究天然混交林林分及主要树种竞争的表达方法,并以甘肃省小陇山林区锐齿槲栎天然混交林为研究对象,采用胸径、树高、冠幅、距离等信息的2期调查数据,对新构建的基于树冠重叠面积的天然混交林林木竞争(CIM)指数进行验证,并选取Spearman系数,对CIM竞争指数和传统竞争指数进行相关性分析。结果表明,采用树冠投影重叠和考虑树高关系确定竞争木的方法能有效避免竞争木多选或漏选; CIM1、CIM2、CIM3能够表达出混交林竞争压力变化,表现出竞争压力越大林木蓄积生长量越小的趋势,将树种混交因子引入竞争指数能够提高蓄积生长量与竞争指数的相关性,更加客观的描述林分中竞争状况。竞争指数与蓄积生长量相关性由大到小排序为CIM2>CIM3>CIM1>Bella竞争指数(CIO)>简单竞争指数(CI)>CIO2>CIO1,表明竞争指数 CIM2对混交林竞争表达更具参考意义。林分中主要树种锐齿槲栎由于相邻木中同树种相邻木增加,导致竞争压力增大。天然混交林中树种组成复杂多样,林木分布不均匀,在竞争指数中引入树种混交因子能够较为真实的反映林木竞争状况。     

杉木-观光木人工混交林林木竞争效应研究

将人工混交林中林木竞争影响空间划分为内、外两圈,采用Hegyi提出的单木竞争指数模型对福建省建瓯东安林场杉木（Cunninghamia lanceolata）一观光木（Tsoongi odenodrum）混交林中林木的种内、种间竞争强度进行分析。结果表明,无论是内圈还是外圈,杉木种内、种间竞争强度均随林木径级的增大而减小;竞争强度顺序为：内圈杉木种内竞争〉内圈杉木观光木种间竞争〉外圈杉木种内竞争〉外圈杉木观光木种间竞争。用模型拟合预测杉木种内、种间的竞争强度,认为杉木在胸径达到36cm之前应采取适当的控制措施。     

Demographic changes and the development of competition in a native Australian eucalypt forest invaded by exotic pines

Data from the second half of a 14-year study of a eucalypt dry sclerophyll forest invaded by Pinus radiata D. Don shows further progress in the development of a mixed eucalypt-pine stand. Earlier (Chilvers and Burdon 1983) it was clear that pines invading from an adjacent commercial plantation were starting to contribute their own progeny to the mixed stand. Initially the recruitment of established pines (> 1 m high) from these seedlings was very slow. However, since 1984, increasing numbers of seedlings have survived beyond their first year, suggesting the possibility of another wave of successful establishment in the near future. The established fraction of the pine population continued to grow rapidly and incurred no mortality. Many of these plants have now emerged through the top of the eucalypt canopy. In contrast, both juvenile and adult eucalypt numbers continued to decline. Despite these demographic and visual changes, comparative analysis of the growth of established pines over two consecutive 7-year periods (1974–1981; 1981–1988) showed that competition between neighbouring plants, especially pines, is beginning to affect individual growth rates. Growth rates of pines having other pines as nearest neighbours declined 40% between 1974–1981 and 1981–1988, while that of pines with eucalypts as nearest neighbours fell by 28%. In pine-pine nearest neighbour (NN) pairs, growth rates were significantly affected by the size (diameter) of the nearest neighbour. In pine-eucalypt NN pairs growth rates of the pines were not affected by the size of the eucalypt, but these were significantly negatively correlated with the inverse of the distance to the eucalypt. More broad-based assessment of 43 pines showed a significant relationship between their growth rates and the summed competitive effect of other pines within a 5-m radius. No such relationship was found between growth rate and the summed competitive effect of eucalypts in the 5-m zone.     

An alternative to Lotka-Volterra competition in coarse-grained environments

Competition in a coarse-grained heterogeneous environment is considered. It is assumed that each individual adjust has position within the macrohabitat so as to try to maximize a utility whose value is a function of position within the habitat and of the population density at that position. Robust qualitative conclusions concerning the resulting population behavior relate niche breadth, niche shift, and resource partitioning to changes in population numbers. In particular populations avoid competition by reducing coarse-grained niche overlap so that overlap is a measure not of the degree of competition but of its absence. Specific models of exploitation or interference competition give specific quantitative results which are summarized. Competition equations are derived from underlying models which imply Lotka-Volterra equations in a fine-grained environment (MacArthur, R., and Levins, R. 1967. Amer. Natur.101, 377–385) but which are nonlinear in per capita growth in a coarse-grained one. The findings are relevant to theoretical studies of competition, limiting similarity, species packing, and the evolution of niche position as well as to practical problems of data analysis and resource management. The extension to predator-prey interactions is outlined.     

落叶松人工林空间结构优化的探讨

基于落叶松(Larix olgensis Henry)人工林固定标准地ML707的每木检尺数据,针对人工林普遍具有混交度低,分布均匀的特点。主要从林分竞争机制和树冠营养空间利用情况等方面提出了利用Hegyi竞争指数、生长空间指数、生长空间竞争指数分析优化林分空间结构,进一步比较了不同择伐强度下林分空间结构变化的大小。研究结果表明：14%左右为林分最优的择伐强度,空间优化达到最佳,优化后林分竞争态势减小,营养空间利用程度增大,林分空间结构得到改善。     

Defoliation and competition effects in a productivity gradient for a semiarid Mediterranean annual grassland community

Grazing and competition are two main factors shaping range plant communities; however, few studies have investigated their interaction. The current study aimed to investigate the effects of defoliation, competition and their interaction on production of annual grasses in semiarid Mediterranean areas. Competition treatments (absence/presence of neighbors) were combined with three defoliation intensities (0%, 30% and 60%) in a complete factorial design. Competition significantly reduced grass biomass. However, the role of competition was eliminated under heavy defoliation or under dry growth conditions. Defoliation showed variable results on final biomass (FB) and cumulative biomass (CB). While heavy defoliation (60% clipping intensity) reduced grass FB down to 80% during the two growing seasons, light defoliation (30%) significantly increased CB. Results showed that competition may limit the direct effect of defoliation on dominant grass species. Further, the relationship between site productivity and competition effect was best explained by a negative linear model. This hypothesized model may suggest that facilitation and competition alternatively affect grassland communities along a productivity gradient. The results suggest that light grazing may sustain or even enhance grassland productivity. The results also indicated the suitability of annual grass species to re-vegetate degraded rangeland in semi-arid climate. Further, optimum grazing practices to conserve biodiversity of Avena grassland may involve moderate stocking rate.     

山东蒙山森林冠层绿度与树干径向生长的关系

森林冠层绿度和树木年轮宽度是描述森林生长过程的重要指标,它们之间存在怎样的关系以及这种关系的稳定性如何目前还没有清晰的回答。森林冠层绿度通过遥感影像计算,在空间上连续,而树木年轮宽度是树木健康的综合指标,样点上具有代表性。森林冠层绿度和树木年轮宽度的关系的研究能增进对森林生长的多角度理解和森林生长状况的尺度转换。在山东蒙山地区采集了4个赤松(Pinus densiflora)林样点的树木年轮样本,获得了树木年轮宽度数据,分析了增强型植被指数(Enhanced Vegetation Index,EVI)与树木年轮宽度的关系。结果显示:1)对于健康森林,4月和6月的冠层绿度与树木年轮宽度存在因果关系;森林不健康时,两者关系较为复杂;2)其他月份冠层绿度与树干径向生长不存在因果关系,而是共同受其他环境因子,如气候因子的驱动;3)弱冠层绿度降低后5年内有显著的径向生长恢复,但是恢复年份少;强冠层绿度降低之前,树干径向生长已经开始降低,之后的5年内有着持续的径向生长降低。这些结果表明森林冠层绿度的降低并不能反映树干径向生长降低的开始,只有健康的森林冠层绿度和年轮宽度有相关关系。冠层绿度的降低对森林健康有强烈的影响,冠层绿度降低导致的径向生长的降低很难恢复。     

Plant competition and exclusion with optimizing individuals

Most models of plant competition represent competition as taking place between species when realistically competition takes place between individuals. We model individual plants as optimally choosing biomass in order to maximize net energy that is directed into reproduction. Competition is for access to light and a plant that grows more biomass adds to the leaf area index, creating negative feedback in the form of more self shading and shading of its neighbors. In each period and for given species densities, simultaneous maximization by all plants yields an equilibrium characterized by optimum biomasses. Between periods the net energies plants obtain are used to update the densities, and if densities change the equilibrium changes in the subsequent period. A steady state is attained when all plants have net energies that just allow for replacement. Four main predictions of the resource-ratio theory of competition are obtained, providing behavioral underpinnings for species level models. However, if individual plant parameters are not identical across species, then the predictions do not follow. The optimization framework yields many other predictions, including how specific leaf areas and resource stress impact biomass and leaf area indices.     

Size bias in traditional analyses of substitutive competition experiments

Summary The objective of this study was to examine the claim that traditional measures of competitive performance in substitutive experiments are biased towards larger plants. Results from a three-year diallele experiment of 6 marsh plant species were analyzed using both Relative Yields (a traditional analysis) and the Relative Efficiency Index (a recently proposed analysis presumed to be size-independent). In adddition, a mechanistic model of competition was used to explore the behavior of both methods of estimating competitive performance.Results from the three-year experiment showed that Relative Yields (RYs) were correlated with the initial sizes of plants for the first two years but not the third. By the third year, RYs were highly correlated with Relative Efficiency Index values (REIs) suggesting that the effects of initial size were eventually overcome. Model results showed that RYs are inherently biased in favor of larger plants during the early phases of competition while REIs are not. Further, model analysis confirmed that the size bias associated with RYs declines with increasing duration of the experiment. It is concluded that current generalizations about the relationship between plant size and competitive ability may be biased by the procedures used to analyze competition experiments.     

Scales of aboveground and below-ground competition in a semi-arid woodland detected from spatial pattern

Abstract. Semi-arid woodlands are two-phase mosaics of canopy and inter-canopy patches. We hypothesized that both aboveground competition (within canopy patches), and below-ground competition (between canopy patches), would be important structuring processes in these communities. We investigated the spatial pattern of trees in a Pinus edulis-Juniperus monosperma woodland in New Mexico using Ripley's K-function. We found strong aggregation of trees at scales of 2 to 4 m, which indicates the scale of canopy patches. Canopy patches were composed of individuals of both species. Crown centers of both species were always less aggregated than stem centers at scales less than canopy patch size, indicating morphological plasticity of competing crowns. In the smallest size classes of both species, aggregation was most intense, and occurred over a larger range of scales; aggregation decreased with increasing size as is consistent with density-dependent mortality from intraspecific competition. Within canopy patches, younger trees were associated with older trees of the other species. At scales larger than canopy patches, younger trees showed repulsion from older conspecifics, indicating below-ground competition. Hence, intraspecific competition was stronger than interspecific competition, probably because the species differ in rooting depth. Woodland dynamics depend on the scale and composition of canopy patches, aggregated seed deposition and facilitation, above- and below-ground competition, and temporal changes in the spatial scale of interactions. This woodland is intermediate in a grassland-forest continuum (a gradient of increasing woody canopy cover) and hence we expected, and were able to detect, the effects of both above- and below-ground competition.     

邻体竞争对植物个体生长速率和死亡概率的影响：基于日本落叶松种群试验的研究

选择 4种有代表性的植物邻体竞争指数 ,分析了竞争对日本落叶松 (L arix leptolepis)试验种群的个体生长速率和死亡概率的影响 ,发现在本试验条件下 ,引入竞争指数不能明显改善对生长速率的预测效果 ,但用竞争指数可以较好地预测个体于 1个月内在竞争压力下死亡的概率 ,从而为构造植物生长模型时处理个体死亡问题提供一个有益的定量化思路