Using approximate Bayesian computation to infer photosynthesis model parameters北大核心CSCD

We developed a method, namely Adaptive Population Monte Carlo Approximate Bayesian Computation (APMC), to estimate the parameters of Farquhar photosynthesis model. Treating the canopy as a big leaf, we applied this method to derive the parameters at canopy scale. Validations against observational data showed that parameters estimated based on the APMC optimization are un-biased for predicting the photosynthesis rate. We conclude that APMC has greater advantages in estimating the model parameters than those of the conventional nonlinear regression models.     

Effects of ozone and aerosol pollution on photosynthesis of poplar leaves

由于经济的快速发展, 中国大部分地区正面临着严峻的复合型大气污染, 其中臭氧和气溶胶是两种主要污染物。已有的研究表明臭氧对叶片的氧化性伤害能够抑制光合作用, 而气溶胶可通过增加散射辐射比例或缓解高温抑制促进光合作用。但复合污染下, 臭氧和气溶胶如何共同调控叶片光合作用, 仍缺乏研究。该研究利用北京及周边地区之间的污染梯度, 选择加杨(Populus × canadensis)作为实验对象, 于2012-2013年生长季期间对叶片光合速率进行连续观测, 并同时监测臭氧浓度(AOT40)、气溶胶光学厚度(AOD)、空气温度和冠层内外光合有效辐射(PAR)等环境因子, 以期探讨大气复合污染下臭氧和气溶胶变化对植物叶片光合作用的影响及相关机制。结果表明: (1)臭氧浓度与空气温度、气溶胶浓度之间均呈显著正相关关系, 但气溶胶浓度与空气温度没有显著相关关系; (2)臭氧浓度增加显著抑制了阳生叶片的光合作用, 但气溶胶浓度上升促进了阳生叶片的光合作用; 臭氧浓度升高对阴生叶片光合作用的影响较小, 但气溶胶浓度上升促进了阴生叶片的光合作用; (3)标准化后的结果显示, 臭氧对阳生叶片光合作用的影响最大, 此时气溶胶的促进作用一定程度上补偿了臭氧浓度上升所带来的抑制效应。对于阴生叶片光合作用而言, 气溶胶则是最重要的影响因素。该研究发现复合污染下阴生叶和阳生叶光合响应不同, 这表明冠层结构可能通过影响阴生叶和阳生叶的比例, 从而对植物生长产生不同影响。该研究对理解大气复合污染如何影响光合作用提供了的机理支持, 同时也表明, 为了维持生态系统生产力及功能, 需要同时控制气溶胶和臭氧污染。     

用光合-蒸散耦合模型模拟冬小麦CO2通量的日变化

根据SPAC理论建立了一个冬小麦光合和蒸散的耦合模型.冬小麦CO2通量包括冠层光合、呼吸和土壤呼吸.冠层光合采用了Farquhar光合作用生化模型,并通过冠层阻力的参数化将光合作用与蒸腾作用耦合起来.用涡度相关方法观测了CO2通量,对模型进行了验证,结果显示模型可以较好地模拟CO2通量日变化过程.对模型的敏感性分析发现日间CO2通量最敏感的参数是初始量子效率.其次,CO2通量对光响应曲线凸度、CO2补偿点、凋萎点和叶面积指数的变化也有着较强的敏感性;夜间CO2通量敏感的参数是最适温度下Rubisco催化能力和暗呼吸参数.     

用光合-蒸散耦合模型模拟冬小麦CO2通量的日变化

根据 SPAC理论建立了一个冬小麦光合和蒸散的耦合模型。冬小麦 CO2 通量包括冠层光合、呼吸和土壤呼吸。冠层光合采用了 Farquhar光合作用生化模型 ,并通过冠层阻力的参数化将光合作用与蒸腾作用耦合起来。用涡度相关方法观测了 CO2通量 ,对模型进行了验证 ,结果显示模型可以较好地模拟 CO2 通量日变化过程。对模型的敏感性分析发现日间 CO2 通量最敏感的参数是初始量子效率。其次 ,CO2 通量对光响应曲线凸度、CO2 补偿点、凋萎点和叶面积指数的变化也有着较强的敏感性 ;夜间 CO2 通量敏感的参数是最适温度下 Rubisco催化能力和暗呼吸参数     

叶绿素荧光主动与被动联合观测应用前景

叶绿素荧光是研究植物光合生理机制、量化植被光合作用时空格局以及准确理解气候变化背景下陆地生态系统生产力的关键。然而, 目前对于叶绿素荧光主动与被动联合观测的研究还较少。该文对比了叶绿素荧光主动观测与被动观测的优缺点, 展示了叶片尺度和冠层尺度主动与被动联合观测的仪器设备组成, 探讨了主动与被动联合观测在探索叶绿体尺度-叶片尺度-冠层尺度能量在光合、荧光以及热耗散中的分配, 阐明叶绿素荧光与总初级生产力的关联机制, 验证星基日光诱导叶绿素荧光, 解译叶绿素荧光光谱形状4个方面的应用前景。综上, 叶绿素荧光的主动与被动联合观测对于揭示各尺度上荧光与光合作用之间的关联机制, 改善全球尺度植被生产力模型至关重要。     

Effect of light interception by non-photosynthetic organs on canopy photosynthetic production

A canopy photosynthesis model was derived on the assumption that the light diminution within a canopy is caused by both leaves and non-photosynthetic organs. The light diminution by leaves and that by non-photosynthetic organs were taken into account separately in the Lambert-Beer equation of light extinction. The light flux density on the leaf surface at each depth was evaluated from the leaf's share of light. The light flux density on the leaf surface thus obtained was incorporated into the Monsi-Saeki model of canopy photosynthesis. The proposed model was applied for estimating gross canopy photosynthesis in a 19-year-oldLarix leptolepis plantation where 38% of the light diminution was due to non-photosynthetic organs. The daily canopy photosynthesis on one summer day calculated using the present model was about 22% less than that calculated by the conventional Monsi-Saeki model, in which light interception by non-photosynthetic organs is neglected. The degree of such reduction in canopy photosynthesis through shading by non-photosynthetic organs was assessed in relation to parameters affecting light extinction, leaf photosynthetic characteristics, and light regime above the canopy.     

Modelling canopy photosynthesis using parameters determined from simple non-destructive measurements

Most models for canopy photosynthesis require a large number of parameters as input which have to be determined by means of direct measurements. Such measurements are usually expensive, time consuming and destructive. The objective of the present study was, therefore, to develop a simple but accurate canopy photosynthesis model based on a minimum number of parameters that can be determined non-destructively. The results from previous studies were used to derive an empirical expression which describes the variation in leaf photosynthetic capacity (P_m) as a function of the light distribution in the canopy. The light distribution itself was calculated with a simple model which assumes only three leaf angle classes (0–30°, 30–60° and 60–90°). The leaf area index was determined indirectly from measurements of direct radiation below the canopy. The result was a model for canopy photosynthesis that requires only a few parameters. These parameters are the leaf photosynthetic capacity at the top of the canopy, the relative frequency of leaves in each of the three leaf angle classes, and the fraction of direct radiation below the canopy. Each of these parameters can be determined by means of simple non-destructive measurements. The model was applied to dense stands of two monocotyledonous species: rice (Oryza sativa L.) and pearl millet (Pennisetum americanum (L.) K. Schum.). The rates of canopy photosynthesis thus calculated were compared to those obtained with a more elaborate reference model. The differences between the values obtained with the two models were small. The present photosynthesis model can, therefore, be considered to be a suitable alternative for the more elaborate model. It was further discussed that, since the model is based on purely non-destructive measurements, it will be particularly useful in cases where it is required to estimate canopy photosynthesis at regular intervals over a length of time or in stands of vegetation that cover large areas of land.     

Modelling the effects of changes in solar radiation on gross primary production in subtropical evergreen needle-leaf plantations

太阳辐射是陆地生态系统碳水循环的能量来源。太阳辐射的变化对植被吸收大气CO₂具有重要影响。该文通过辐射观测数据建立散射辐射比例与晴空指数的关系, 结合生态过程模型(BEPS)和通量观测数据, 模拟分析了太阳辐射变化对千烟洲常绿针叶林总初级生产力(GPP)的影响。研究结果表明: 千烟洲森林生态系统的阴叶对年GPP总量的贡献达67%, 太阳辐射变化对阴叶光合作用的影响决定了冠层GPP的变化; 太阳辐射强度和分布的年际差异导致年GPP对太阳辐射变化的响应不同, 2003、2004和2005年太阳辐射分别变化-5.44%、-1.83%和6.26%, 可使千烟洲生态系统当年GPP总量达到最大值; 在季节上, 太阳辐射的增加会导致5-6月GPP上升, 7-9月GPP下降, 使年GPP变化程度降低; 在天尺度上, 晴空指数在0.43时, 太阳辐射变化对GPP的影响最小。     

用光合-蒸散耦合模型模拟冬小麦水热通量的日变化

从SPAC理论出发,建立了一个冬小麦光合和蒸散的耦合模型．感热通量和潜热通量采用Shuttleworth-Wallace的双层模型计算,并通过冠层阻力的参数化,将光合作用与蒸腾作用耦合起来．用涡度相关方法,观测了感热通量和潜热通量,对模型进行了验证．结果表明,模拟值与观测值比较一致,模型可以很好地模拟感热通量和潜热通量的日变化过程．对模型的敏感性分析发现,冬小麦蒸腾比较敏感的参数有凋萎点、气孔导度参数、叶对红外辐射的反射率和光响应曲线凸度;土壤蒸发只对土壤阻力参数的敏感性较强．本模型对水热通量与环境因子作用过程的理论研究和指导农田的灌溉制度等有一定的意义．     

Exploring the spatial distribution of light interception and photosynthesis of canopies by means of a functional-structural plant model

Background and Aims

At present most process-based models and the majority of three-dimensional models include simplifications of plant architecture that can compromise the accuracy of light interception simulations and, accordingly, canopy photosynthesis. The aim of this paper is to analyse canopy heterogeneity of an explicitly described tomato canopy in relation to temporal dynamics of horizontal and vertical light distribution and photosynthesis under direct- and diffuse-light conditions.

Methods

Detailed measurements of canopy architecture, light interception and leaf photosynthesis were carried out on a tomato crop. These data were used for the development and calibration of a functional–structural tomato model. The model consisted of an architectural static virtual plant coupled with a nested radiosity model for light calculations and a leaf photosynthesis module. Different scenarios of horizontal and vertical distribution of light interception, incident light and photosynthesis were investigated under diffuse and direct light conditions.

Key Results

Simulated light interception showed a good correspondence to the measured values. Explicitly described leaf angles resulted in higher light interception in the middle of the plant canopy compared with fixed and ellipsoidal leaf-angle distribution models, although the total light interception remained the same. The fraction of light intercepted at a north–south orientation of rows differed from east–west orientation by 10 % on winter and 23 % on summer days. The horizontal distribution of photosynthesis differed significantly between the top, middle and lower canopy layer. Taking into account the vertical variation of leaf photosynthetic parameters in the canopy, led to approx. 8 % increase on simulated canopy photosynthesis.

Conclusions

Leaf angles of heterogeneous canopies should be explicitly described as they have a big impact both on light distribution and photosynthesis. Especially, the vertical variation of photosynthesis in canopy is such that the experimental approach of photosynthesis measurements for model parameterization should be revised.     

Simple scaling of photosynthesis from leaves to canopies without the errors of big-leaf models

In big-leaf models of canopy photosynthesis, the Rubisco activity per unit ground area is taken as the sum of activities per unit leaf area within the canopy, and electron transport capacity is similarly summed. Such models overestimate rates of photosynthesis and require empirical curvature factors in the response to irradiance. We show that, with any distribution of leaf nitrogen within the canopy (including optimal), the required curvature factors are not constant but vary with canopy leaf area index and leaf nitrogen content. We further show that the underlying reason is the difference between the time-averaged and instantaneous distributions of absorbed irradiance, caused by penetration of sunflecks and the range of leaf angles in canopies. These errors are avoided in models that treat the canopy in terms of a number of layers – the multi-layer models. We present an alternative to the multi-layer model: by separately integrating the sunlit and shaded leaf fractions of the canopy, a single layered sun/shade model is obtained, which is as accurate and simpler. The model is a scaled version of a leaf model as distinct from an integrative approach.     

A comparison of measured and calculated net community CO2 exchange: Scaling from leaves to communities

Leaf net photosynthesis is crucial for detecting the mechanism of photosynthesis, whereas community net photosynthesis is useful for understanding the photosynthetic capacity of communities and its relationship with environmental factors. In particular, we need to scale up eco-physiological models from leaf scale to canopy level to study carbon cycling at regional or global scale. We hypothesized that accumulated leaf net photosynthetic rate (P_c) at community scale, i.e., calculated based on leaf net photosynthetic rate (P_n) and leaf area index (LAI), equals to measured net community CO₂ exchange (NCE). The purpose of this study is to verify this hypothesis. Our field study was carried out in Duolun, Nei Mongol, China, where we constructed single-species communities by sowing Medicago sativa ‘Aohan’ seeds in three plots (3 m × 5 m) on May 30, 2012. On August 16, 2014, P_n of five healthy leaves of M. sativa ‘Aohan’ in each plot were measured with a LI-6400 portable photosynthesis system at 10:00, and net ecosystem CO₂ exchange (NEE) in each plot was measured simultaneously with a LI-8100 system connected with a assimilation chamber (0.5 m × 0.5 m × 0.5 m). P_c was calculated based on P_n, number of leaves (n), LAI percentage of healthy leaves (r) and percentage of received effective light by leaves (m). NCE was derived from NEE and ecosystem respiration rate (R_eco). P_c was 3.52 μmol CO₂·m^-2·s^-1, and very close to NCE (3.56 μmol CO₂·m^-2·s^-1), suggesting that leaf-scale photosynthesis may accurately predict community-scale photosynthesis. However, our method could not separate community respiration from soil respiration, and future studies, should be designed to counteract this effect. Scaling up from leaf photosynthesis to community photosynthesis should also consider vertical structure of communities and nonlinear responses of leaf photosynthesis to changes in light quantum.     

Modeling study on photosynthetic-light response curves of a C4 plant,maize

Aims A light response curve can reflect a plant’s ability to utilize light, which is also a key tool in determining the relationship between photosynthetic capacity and environmental factors; however the model accuracies concerning the light response curve remain elusive. The objectives of this study were to compare and assess the model accuracies related to a light response curve and the effects of drought. Methods A field rain shelter was used to control the soil water conditions. To obtain photosynthesis parameters from the light response curve and the drought effects, the relevant models (including the rectangular model, non-rectangular hyperbolic model, modified rectangular hyperbolic model, exponential model, quadratic function model, and a newly modified model) were applied to fit the light response curves. The validity of each model was tested by analyzing the differences between the fitted values obtained by the models and the measured values. Important findings The newly modified model has been proved to performing relatively better in accurately describing the light response curve patterns, and credibly obtaining the crucial photosynthetic parameters such as the maximum net photosynthetic rate, light saturation point, light compensation point, and dark respiration rate, especially under high radiation conditions.     

Shrub encroachment effect on the evapotranspiration and its component—A numerical simulation study of a shrub encroachment grassland in Nei Mongol,China

Aims Shrub encroachment is a common global change phenomenon occurring in arid and semi-arid regions. Due to the difficulty of partitioning evapotranspiration into shrub plants, grass plants and soil in the field, there are few studies focusing on shrub encroachment effect on the evapotranspiration and its component in China. This study aims to illustrate shrub encroachment effect on evapotranspiration by the numerical modeling method. Methods A two-source model was applied and calibrated with the measured evapotranspiration (ET) by the Bowen ratio system to simulate evapotranspiration and its component in a shrub encroachment grassland in Nei Mongol, China. Based on the calibrated model and previous shrub encroachment investigation, we set three scenarios of shrub encroachment characterized by relative shrub coverage of 5%, 15% and 30%, respectively, and quantified their effects caused by shrub encroachment through localized and calibrated two-source model.Important findings The two-source model can well reconstruct the evapotranspiration characteristics of a shrub encroachment grassland. Sensitivity analysis of the model shows that errors for the input variables and parameters have small influence on the result of partitioning evapotranspiration. The result shows that shrub encroachment has relatively small influence on the total amount of ET, but it has clear influence on the proportion of the components of evapotranspiration (E/ET). With shrub coverage increasing from 5% to 15% and then 30%, the evapotranspiration decreased from 182.97 to 180.38 and 176.72 W·m^-2, decreasing amplitude values of 0.34% and 0.44%, respectively. On average, E/ET rises from 52.9% to 53.9% and 55.5%, increasing amplitude values to 2.04% and 3.25%. Data analysis indicates that shrub encroachment results in smaller soil moisture changes, but clear changes of ecosystem structure (decreasing ecosystem leaf area index while increasing vegetation height) which lead to the decrease of transpiration fraction through decreasing canopy conductance. The research highlights that, with the shrub encroachment, more water will be consumed as soil evaporation which is often regarded as invalid part of evapotranspiration and thus resulting in the decrease of water use efficiency.     

Development of the Monsi-Saeki theory on canopy structure and function

BACKGROUND AND AIMS: Monsi and Saeki (1953) published the first mathematical model of canopy photosynthesis that was based on the light attenuation within a canopy and a light response of leaf photosynthesis. This paper reviews the evolution and development of their theory. SCOPE: Monsi and Saeki showed that under full light conditions, canopy photosynthesis is maximized at a high leaf area index (LAI, total leaf area per unit ground area) with vertically inclined leaves, while under low light conditions, it is at a low LAI with horizontal leaves. They suggested that actual plants develop a stand structure to maximize canopy photosynthesis. Combination of the Monsi-Saeki model with the cost-benefit hypothesis in resource use led to a new canopy photosynthesis model, where leaf nitrogen distribution and associated photosynthetic capacity were taken into account. The gradient of leaf nitrogen in a canopy was shown to be a direct response to the gradient of light. This response enables plants to use light and nitrogen efficiently, two resources whose supply is limited in the natural environment. CONCLUSION: The canopy photosynthesis model stimulated studies to scale-up from chloroplast biochemistry to canopy carbon gain and to analyse the resource-use strategy of species and individuals growing at different light and nitrogen availabilities. Canopy photosynthesis models are useful to analyse the size structure of populations in plant communities and to predict the structure and function of future terrestrial ecosystems.     

杉木人工林单叶至冠层光合作用的扩展与模拟研究

根据野外条件下杉木（Ｃｕｎｎｉｎｇｈａｍｉａ ｌａｎｃｅｏｌａｔａ（Ｌａｍｂ．）Ｈｏｏｋ．）针叶光合作用的测定结果。考虑到光合作用对光的非线性响应特性,及其与叶所处的实际冠层环境变量和冠层的空气动力学特性的相关,以筒化的林冠辐射传输模型为基础,结合不同部位和年龄针叶的光响应曲线,实现了叶室测量结果向冠层环境的调整,并进行了冠层光合作用模拟的初步研究。经过调整后的冠层光合作用平均比未经过调整的值高１     

Leaf photosynthetic light response: a mechanistic model for scaling photosynthesis to leaves and canopies

1. The response of photosynthesis to radiation is an often-studied but poorly understood process, represented empirically in most photosynthesis models. However, in scaling photosynthesis from leaf to canopy, predictions of canopy photosynthesis are very sensitive to parameters describing the response of leaves to Photosynthetic Photon Flux Density (PPFD).
2. In this study, a mechanistic, yet still simple, approach is presented that models the degree of light saturation in leaves explicitly, assuming a heterogeneous environment of PPFD and chlorophyll.
3. Possible mechanisms determining the ratio of chlorophyll to nitrogen are considered, including a direct dependence on PPFD, a mechanism involving the red/far-red ratio of light in the canopy, and an approach based upon maximizing photosynthesis.
4. Comparison of model predictions with two data sets of light, nitrogen and chlorophyll from canopies of Populus and Corylus suggests that the red/far-red mechanism is the most realistic. The data also show that the trees studied do not always optimize their nitrogen partitioning to maximize photosynthetic yield.
5. We then apply the model to the data sets, to predict the shape of light response curves of leaves within canopies and assess the applicability of simple scaling schemes, in which full acclimation of photosynthesis to PPFD justifies the use of big-leaf models. We conclude that, at least for the data used, basic assumptions of such schemes do not hold.     

Plant competition for light analyzed with a multispecies canopy model

Summary Competition for light among species in a mixed canopy can be assessed quantitatively by a simulation model which evaluates the importance of different morphological and photosynthetic characteristics of each species. A model was developed that simulates how the foliage of all species attenuate radiation in the canopy and how much radiation is received by foliage of each species. The model can account for different kinds of foliage (leaf blades, stems, etc.) for each species. The photosynthesis and transpiration for sunlit and shaded foliage of each species is also computed for different layers in the canopy. The model is an extension of previously described single-species canopy photosynthesis simulation models. Model predictions of the fraction of foliage sunlit and interception of light by sunlit and shaded foliage for monoculture and mixed canopies of wheat (Triticum aestivum) and wild oat (Avena fatua) in the field compared very well with measured values. The model was used to calculate light interception and canopy photosynthesis for both species of wheat/wild oat mixtures grown under normal solar and enhanced ultraviolet-B (290–320 nm) radiation (UV-B) in a glasshouse experiment with no root competition. In these experiments, measurements showed that the mixtures receiving enhanced UV-B radiation had a greater proportion of the total foliage area composed of wheat compared to mixtures in the control treatments. The difference in species foliage area and its position in the canopy resulted in a calculated increase in the portion of total canopy radiation interception and photosynthesis by wheat. This, in turn, is consistent with greater canopy biomass of wheat reported in canopies irradiated with supplemental UV-B.     

Exploring the influence of soil types underneath the canopy in winter wheat leaf area index remote estimating

Aims Remote sensing is an effective and nondestructive way to retrieve leaf area index (LAI) from plot, regional and global range. Soil background is one of the confounding factors limiting remotely estimating LAI. And soil type contains a large proportion of soil background information, which can influence the optical properties of vegetation canopy and soil. However, our knowledge on the effects stemmed from soil types underneath the canopy on LAI remote estimating have been in shortage. Thus, this study aims to explore the influences of soil types underneath the canopy on winter wheat LAI remote estimating. Methods We analyzed the sensitivity variation of eight spectral indices, named normalized difference vegetation index (NDVI), modified soil-adjusted vegetation index (MSAVI), modified chlorophyll absorption ratio index 2 (MCARI₂), red edge inflection point (REIP), red edge amplitude (Dr), red edge area (SDr), red edge symmetry (RES), normalized difference spectral index (NDSI), to LAI in different soil types, and then we identified some spectral intervals or parameters that were insensitive to soil type variations underneath the canopy. We also compared the accuracy of two commonly used regression models, partial least squares regression (PLSR) and random forest regression (RFR), in estimating LAI for different soil types. We also explored the problems arising from applying the regression model developed in single soil type area to complex soil types area in retrieving LAI. Important findings This paper demonstrates the effects of soil types underneath the canopy on LAI retrieving. 1) The sensitivity of spectral indices to LAI is significantly different due to the soil type variation, but REIP has the least effects from soil type variation among the eight spectral indices. Meanwhile, the band selection algorithm of lambda-by-lambda not only chooses the most sensitive spectral interval for LAI, but also provides a feasible way to construct the spectral index that exhibits strong resistances to the effects of soil types underneath the canopy. 2) The accuracy of LAI estimation by regression models differs under soil type considered or not. So we suggest that in small scale researches, especially in a field scale, the ability of regression models in explaining variables is the priority consideration, while the PLSR is superior to RFR in this respect. Under the premise of unknown priori knowledge of land surfaces, the RFR is more suitable for retrieving LAI than PLSR, but land surface priori knowledge is still necessary. These findings provide the theoretical basis and methods for developing remotely sensing estimating LAI models adapted to various land surfaces. Further analysis is needed in applying the findings in more crop types, cultivars and growth stages.     

树冠结构对典型阔叶红松林生产力的影响

阔叶红松(Pinus koraiensis)林是东北东部山区的地带性森林植被, 阐明其生产力的影响因素, 对于理解温带森林生产力维持机制具有重要意义。该研究依托小兴安岭典型阔叶红松林9 hm²动态监测样地, 基于2005和2015年的30 m × 30 m样方内所有胸径>6.5 cm的木本植物的调查数据, 计算各样方的树冠结构复杂性、物种多样性和林分胸高断面积, 结合各样方的地形和土壤理化性质数据, 拟合结构方程模型, 定量分析影响典型阔叶红松林生产力的直接和间接因素。研究结果显示: 树冠结构复杂性和物种多样性与生产力显著正相关, 且树冠结构复杂性对生产力的影响显著高于物种多样性; 树冠结构复杂性对生产力的作用分为树冠垂直分层和树冠可塑性, 其中树冠垂直分层是树冠结构复杂性影响阔叶红松林生产力的主要因素, 而树冠可塑性无显著影响; 林分胸高断面积与生产力显著正相关, 其解释权重仅次于树冠结构复杂性, 树冠结构复杂性与物种多样性均通过影响林分胸高断面积对阔叶红松林生产力产生间接影响; 考虑不同树冠结构复杂性时, 坡度和土壤全磷含量代表的环境因素在调节生产力上发挥的作用存在差异, 移除树冠垂直分层的作用后两者与生产力呈显著的负相关关系。综上可知, 在典型阔叶红松林中, 树冠结构复杂性比物种多样性更有效地解释了生产力的变化, 同时不可忽视其他生物和非生物因素对生产力的作用。