Spatial and temporal variations in photosynthetic capacity of a temperate deciduous-evergreen forest

Key message

The understory evergreen trees showed maximal photosynthetic capacity in winter, while the overstory deciduous trees showed this capacity in spring. The time lag in productive ecophysiologically active periods between deciduous overstory and evergreen understory trees in a common temperate forest was clearly related to the amount of overstory foliage.

Abstract

In temperate forests, where deciduous canopy trees and evergreen understory trees coexist, understory trees experience great variation in incident radiation corresponding to canopy dynamics represented by leaf-fall and leaf-out. It is generally thought that changes in the light environment affect understory plants’ ecophysiological traits. Thus, to project and estimate annual energy, water, and carbon exchange between forests and the atmosphere, it is necessary to investigate seasonal variation in the ecophysiological activities of both evergreen trees in the understory and deciduous trees that make up the canopy/overstory. We conducted leaf-scale gas-exchange measurements and nitrogen content analyses for six tree species along their heights throughout a complete year. Photosynthetic capacity as represented by the maximum carboxylation rate (V _cmax25) and photosynthetic nitrogen use efficiency (PNUE) of deciduous canopy trees peaked immediately after leaf-out in late May, declined and stabilised during the mid-growing season, and drastically decreased just before leaf-fall. On the other hand, the timing of lowest V _cmax25 and PNUE for evergreen understory trees coincided with that of the highest values for canopy trees. Furthermore, understory trees’ highest values appeared just before canopy tree leaf-out, when incident radiation in the understory was highest. This implies that failing to consider seasonal variation in leaf ecophysiological traits for both canopy and understory trees could lead to serious errors in estimating ecosystem productivity and energy balance for temperate forests.

     

DIFFERENCES IN LEAF STRUCTURE,CHLOROPHYLL, AND NUTRIENTS FOR THE UNDERSTORY TREE ASIMINA TRILOBA

To investigate differences in leaf structure, chlorophyll and nutrients on terminal branches of the understory tree Asimina triloba, the first (proximal) and the last (distal) leaves to develop in the spring were compared. Proximal leaf expansion was completed before the overstory canopy was fully closed but distal leaf expansion occurred during and after the development of the overstory canopy. Fully expanded proximal leaves were 76% smaller in area, were 18% thicker and had 36% more stomates per m of leaf area when compared to distal leaves. In addition, maximum stomatal conductance to water vapor was greater (150 vs. 120 mmol m^−-2s^−-1) and the minimum PPFD required for maximum conductance was higher (200 vs. 150 μmol m^−-2s^−-1) for the proximal leaves. Chlorophyll content was also greater for proximal leaves, but nitrogen and phosphorus contents were lower throughout the entire summer. Seasonal measurements indicated an increase in chlorophyll a content and reductions in nitrogen content throughout the summer growth period for leaves from both positions. The results suggest that distal and proximal leaves differed physiologically and that the measured differences were related to the changing irradiance environment during leaf development. The time of leaf expansion, as indicated by leaf position on the branch, may be an important consideration when examining the water and photosynthetic relations of understory trees.     

浙江天童常绿阔叶林不同演替阶段木本植物的水力结构特征

水力结构是植物应对环境形成的与水分运输相关的形态策略.探索不同演替阶段和群落不同高度层植物的水力结构特征, 有助于理解植物的水分运输和利用策略.该研究以浙江天童常绿阔叶林演替前中后期群落的上层木(占据林冠层的树种)和下层木(灌木层物种)为对象, 测定了演替共有种(至少存在于两个演替阶段的物种)和更替种(仅存在于某一演替阶段的物种)的枝边材比导率,叶比导率和胡伯尔值, 以及边材疏导面积,末端枝总叶面积和枝条水势, 分析植物水力结构在群落上层木和下层木间以及在演替阶段间的差异, 及其与枝叶性状的相关关系.结果显示: (1)上层木植物边材比导率和叶比导率显著高于下层木植物(p < 0.05); (2)上层木和下层木的边材比导率与叶比导率在演替阶段间均无显著差异(p > 0.05); 上层木的胡伯尔值在演替阶段间无显著差异, 下层木的胡伯尔值随演替显著下降(p < 0.05); (3)上层木共有种仅边材比导率随演替进行显著降低(p < 0.05), 更替种的3个水力结构参数在演替阶段间无显著差异; 下层木共有种水力结构参数在演替阶段间无明显差异, 更替种仅胡伯尔值随演替减小(p < 0.05); (4)植物边材比导率与枝疏导面积和末端枝所支撑的总叶面积显著正相关(p < 0.01), 胡伯尔值与枝条水势及末端枝总叶面积显著负相关(p < 0.01).以上结果表明: 天童常绿阔叶林演替各阶段上层木比下层木具有更大的输水能力和效率; 随着演替进行, 上层木与下层木的共有种和更替种边材比导率的相反变化表明上层木水力结构的变化可能由微生境变化引起, 而下层木水力结构特征的变化可能由物种更替造成.     

Hydraulic architecture of evergreen broad-leaved woody plants at different successional stages in Tiantong National Forest Park,Zhejiang Province,China

Aims Hydraulic architecture is a morphological strategy in plants to transport water in coping with environmental conditions. Change of hydraulic architecture for plants occupying different canopy layers within community and for the same plant at different successional stages reflect existence and adaptation in plant's water transportation strategies. The objective of this study was to examine how hydraulic architecture varies with canopy layers within a community and with forest succession.Methods The study site is located in Tiantong National Forest Park, Zhejiang Province, China. Hydraulic architectural traits studied include sapwood-specific hydraulic conductivity, leaf-specific hydraulic conductivity, Huber value, sapwood channel area of twigs, total leaf area per terminal twig, and water potential of twigs. We measured those traits for species that occur in multiple successional stages (we called it "overlapping species") and for species that occur only in one successional stage (we called it "turnover species") along a successional series of evergreen broadleaved forests. For a given species, we sampled both overstory and understory trees. Hydraulic architectural traits between overstory and understory trees in the same community and at successional stages were compared. Pearson correlation was used to exam the relationship between hydraulic architectural traits and the twig/leaf traits.Important findings Sapwood-specific hydraulic conductivities and leaf-specific hydraulic conductivities were significantly higher in overstory trees than those in understory trees, but did not significantly differ from successional stages. Huber value decreased significantly for understory trees, but did not change for overstory trees through forest successional stages. For overstory trees, a trend of decreasing sapwood-specific hydraulic conductivity was observed for overlapping species but not for turnover species with successional stages. In contrast, for understory trees, a trend of decreasing Huber values was observed for turner species but not for overlapping species with successional stages. Across tree species, sapwood-specific hydraulic conductivity was positively correlated with sapwood channel area and total leaf area per terminal twig size. Huber value was negatively correlated to water potential of twigs and total leaf area per terminal twig size. These results suggest that water transportation capacity and efficiency are higher in overstory trees than in understory trees across successional stages in evergreen broadleaved forests in Tiantong region. The contrasting trends of sapwood-specific hydraulic conductivity between overlapping species and turnover species indicate that shift of microenvironment conditions might lead to changes of hydraulic architecture in overstory trees, whereas species replacement might result in changes of hydraulic architecture in understory trees.     

INTERACTIONS BETWEEN CROWN STRUCTURE AND LIGHT ENVIRONMENT IN FIVE RAIN FOREST PIPER SPECIES

Measurements of light variation among leaves within crowns of five Piper species were compared with estimates of spatial variation in light within understory, forest edge, and clearing habitats to estimate the extent to which crown structure contributes to variation in leaf light environment. Daily photon flux density (PFD) varied greatly within and among crowns. Coefficients of variation for daily PFD among sensors within a single crown ranged from 26 to 79%. Within a single crown located in a clearing, the range in daily PFD among leaves was nearly as great as the range over the entire sample of plants. In the understory, localized sunfleck activity contributed to a high degree of spatial variation in instantaneous and total PFD among leaves within individual crowns. Much of the microsite variation in sunfleck activity, however, reflected environmental conditions within the understory habitat. Within an array of sensors placed next to Piper crowns in the understory, correlations were poor for light sensors spaced only 0.2 m apart, and only 8% of the variance in light readings was explained by measurements made 0.5 m away. In the clearing habitats, microsite heterogeneity among leaves was more strongly influenced by leaf positions within crowns and leaf angles than by spatial heterogeneity within the habitat.     

林下层植被对上层乔木的影响研究综述

植物间交互作用在植物群落和生态系统的组成、结构、功能等方面发挥着重要作用.在过去的森林生态系统研究中,更多地关注上层乔木之间的相互作用或乔木层对下层植被的影响,较少研究林下层植被对上层乔木生理生态和生长的影响.本文综述了去除林下层植被对土壤理化性质、土壤动物区系、凋落物分解及上层乔木生理生态和生长的影响,讨论了外界干扰对林下植被-乔木层竞争关系的影响,提出林下植被对上层乔木影响的生理生态学机理概念模型.研究区域、乔木林龄、地力条件、林下植物种类是影响林下层植被-乔木层竞争关系的重要因素.     

Influence of elevated CO2 on canopy development and red:far-red ratios in two-storied stands ofRicinus communis

Vertical structure of plant stands and canopies may change under conditions of elevated CO₂ due to differential responses of overstory and understory plants or plant parts. In the long term, seedling recruitment, competition, and thus population or community structure may be affected. Aside from the possible differential direct effects of elevated CO₂ on photosynthesis and growth, both the quantity and quality of the light below the overstory canopy could be indirectly affected by CO₂-induced changes in overstory leaf area index (LAI) and/or changes in overstory leaf quality. In order to explore such possible interactions, we compared canopy leaf area development, canopy light extinction and the quality of light beneath overstory leaves of two-storied monospecific stands ofRicinus communis exposed to ambient (340 μl l⁻¹) and elevated (610 μl l⁻¹) CO₂. Plants in each stand were grown in a common soil as closed “artificial ecosystems” with a ground area of 6.7 m². LAI of overstory plants in all ecosystems more than doubled during the experiment but was not different between CO₂ treatments at the end. As a consequence, extinction of photosynthetically active radiation (PAR) was also not altered. However, under elevated CO₂ the red to far-red ratio (R:FR) measured beneath overstory leaves was 10% lower than in ecosystems treated with ambient CO₂. This reduction was associated with increased thickness of palisade layers of overstory leaves and appears to be a plausible explanation for the specific enhancement of stem elongation of understory plants (without a corresponding biomass response) under elevated CO₂. CO₂ enrichment led to increased biomass of overstory plants (mainly stem biomass) but had no effect on understory biomass. The results of this study raise the possibility of an important indirect effect of elevated CO₂ at the stand-level. We suggest that, under elevated CO₂, reductions in the R:FR ratio beneath overstory canopies may affect understory plant development independently of the effects of PAR extinction.     

海岸带复合农林业系统植物种群光环境特征研究

本文对三种银杏-农作物复合模式下植物种群对光合有效辐射的削弱和截获进行了模型化分析,并探讨了不同模式下光照强度的时空分布规律。结果表明:银杏果用-叶用-豆类作物复合模式具有最好的复合光效益,光能截获率可达92%;PAR在植冠层的削弱遵循Beer-Lambert定律;植物种群的叶面积、地上部分生物量、光吸收的关系可以用Y=ax^b数学模型描述。同时,果用、材用银杏冠层有较大的透光性和光强变异系数,可以作为银杏-农作物初期经营的上层树种,但要注意冠形的调控;光强的时间变化受冠层条件和太阳高度角的双重影响。     

A tropical epiphytic orchid uses a low‐light interception strategy in a spatially heterogeneous light environment

Light is considered a non‐limiting factor for vascular epiphytes. Nevertheless, an epiphyte's access to light may be limited by phorophyte shading and the spatio‐temporal environmental patchiness characteristic of epiphytic habitats. We assessed the extent to which potential light interception in Rodriguezia granadensis, an epiphytic orchid, is determined by individual factors (plant size traits and leaf traits), or environmental heterogeneity (light patchiness) within the crown of the phorophyte, or both. We studied 104 adult plants growing on Psidium guajava trees in two habitats with contrasting canopy cover: a dry tropical forest edge, and isolated trees in a pasture. We recorded the number of leaves and the leaf area, the leaf position angles, and the potential exposure of the leaf surface to direct irradiance (silhouette area of the leaf blade), and the potential irradiance incident on each plant. We found the epiphytes experience a highly heterogeneous light environment in the crowns of P. guajava. Nonetheless, R. granadensis plants displayed a common light interception strategy typical of low‐light environments, resembling terrestrial, forest understory plants. Potential exposure of the total leaf surface to direct irradiance correlated positively with plant size and within‐plant variation in leaf orientation. In many‐leaved individuals, within‐plant variation in leaf angles produced complementary leaf positions that enhanced potential light interception. This light interception strategy suggests that, in contrast to current wisdom, enhancing light capture is important for vascular epiphytes in canopies with high spatio‐temporal heterogeneity in light environments.     

辽东山区次生林不同大小林窗光照特征比较

以辽东山区天然次生林中3种不同大小林窗(G₁,670 m²;G₂,290 m²和G₃,90 m²)为对象,通过对林窗内光强进行连续观测,比较光量子通量密度(PPFD)的时空分布.结果表明：3种林窗的PPFD日变化均呈现北高南低,且面积越大,PPFD高值区范围越广,异质性越明显;3种林窗的PPFD月变化规律为：林窗内各方位PPFD最大值集中在生长季初期(4—5月),最小值出现的月份则有所差异;3种林窗东部和西部的PPFD出现极值的时间基本一致,且春季光强均明显高于夏、秋季(P<0.05);G₁、G₂、G₃中心点的月平均PPFD分别占全光照的66.59%、49.05%和30.37%,在生长旺盛期,中心点光强分别是林内的37.8倍、27.9倍和10.3倍.受林窗面积不同,以及地形、边缘木高度(林窗形状)等因素的影响,不同大小林窗接收的光强及其分布格局不同,这是导致林窗内更新格局、物种组成发生变化的关键因素.     

Response to CO2 enrichment of understory vegetation in the shade of forests

Responses of forest ecosystems to increased atmospheric CO₂ concentration have been studied in few free‐air CO₂ enrichment (FACE) experiments during last two decades. Most studies focused principally on the overstory trees with little attention given to understory vegetation. Despite its small contribution to total productivity of an ecosystem, understory vegetation plays an important role in predicting successional dynamics and future plant community composition. Thus, the response of understory vegetation in Pinus taeda plantation at the Duke Forest FACE site after 15–17 years of exposure to elevated CO₂, 6–13 of which with nitrogen (N) amendment, was examined. Aboveground biomass and density of the understory decreased across all treatments with increasing overstory leaf area index (LAI). However, the CO₂ and N treatments had no effect on aboveground biomass, tree density, community composition, and the fraction of shade‐tolerant species. The increases of overstory LAI (~28%) under elevated CO₂ resulted in a reduction of light available to the understory (~18%) sufficient to nullify the expected growth‐enhancing effect of elevated CO₂ on understory vegetation.     

Seasonal changes in light and temperature affect the balance between light harvesting and light utilisation components of photosynthesis in an evergreen understory shrub

In a temperate climate, evergreen species in the understory are exposed to large changes in photosynthetic photon flux density (PPFD) and temperature over the year. We determined the photosynthetic traits of leaves of an evergreen understory shrub Aucuba japonica at three sites at monthly intervals: understorys of a deciduous forest; an evergreen forest; and a gap in a mixed forest. This set up enabled us to separate the effects of seasonal change in PPFD and temperature on photosynthetic acclimation under natural conditions. The effects of PPFD and temperature were analysed by simple and multiple regression analyses. The amounts of light utilisation components (LU), represented by nitrogen and rubisco contents per area, were higher in winter, when temperature was low and PPFD was high. The LU relative to the amount of light harvesting components (LH), represented by chlorophyll a/b and rubisco/chlorophyll ratios, and the inverse of chlorophyll/nitrogen ratio were also higher in winter. We quantified the effects of PPFD and temperature on the LU and LH components. Across sites PPFD had stronger effects than air temperature, while within a site temperature had stronger effects on photosynthetic acclimation. We concluded that the photosynthetic apparatus is strongly affected by the prevailing PPFD at the time of leaf development. Within a given light regime, however, plants acclimated by increasing LU relative to LH primarily in response to temperature and to a lesser extent to PPFD.     

林火干扰对北方针叶林林下植被的影响

林下植被在北方针叶林植被群落中的物种多样性最高, 且具有较高的生物量周转率和地上部分净初级生产力, 对北方针叶林生态系统功能起着重要作用。火干扰是决定北方针叶林林下植被结构与功能的一个重要景观过程。该文综述了火干扰是如何通过与地形、火前林冠组成的交互作用而影响环境资源和林下植被的。最近的研究表明: 林下植被能够影响火后树木更新苗的定植、重建速率及森林演替轨迹; 林下植被还会通过影响元素的生物地球化学过程(凋落物降解和养分循环)影响林下环境资源的数量与异质性。因此, 研究火后初期北方针叶林林下植被的动态变化, 对于物种多样性保护和森林管理具有重要意义。     

Deciduousness Influences the Understory Community in a Semideciduous Tropical Forest

We investigated how deciduousness of overstory tree species influences the community structure and species composition in the understory. The results suggest that deciduous overstory trees have positive effects on light‐demanding species, and that the processes underlying such effects may involve reduced competition for light or facilitation through increased water availability.     

Response of overstory and understory vegetation 37 years after prescribed burning in an aspen-dominated forest in northern Minnesota,USA – A case study

Many studies have examined short-term changes in understory vegetation following prescribed burning. However, knowledge concerning longer term effects on both forest understory and overstory vegetation is lacking. This investigation was initiated to examine changes in understory (herbaceous and shrub) and overstory species composition almost four decades after logging and prescribed burning at the Pike Bay Experimental Forest in Minnesota. The experiment was established in 1964 with a randomized block design with four treatments: control (c); burned in spring 1967 (S0); burned in spring 1967 + repeat burn spring 1969 (S2); and burned in spring 1967 + repeat burn fall 1970 (F4). Overstory and understory species diversity indices and richness varied within and among treatments but were not strongly or consistently affected by the treatments. Multivariate analyses (multi-response block permutation procedures and non-metric multidimensional scaling) reveal some lingering effects of burning intensity and seasonal variation as well as some compositional differentiation among treatments, but only in the herb layer. In this environment, the effects of two repeated burnings (fire) have essentially disappeared for overstory and understory species diversity and community composition and have failed to convert an aspen-dominated stand to a coniferous stand (an original goal of the study).     

Ground and remote sensing-based measurements of leaf area index in a transitional forest and seasonal flooded forest in Brazil

Leaf area index (LAI) is a key driver of forest productivity and evapotranspiration; however, it is a difficult and labor-intensive variable to measure, making its measurement impractical for large-scale and long-term studies of tropical forest structure and function. In contrast, satellite estimates of LAI have shown promise for large-scale and long-term studies, but their performance has been equivocal and the biases are not well known. We measured total, overstory, and understory LAI of an Amazon-savanna transitional forest (ASTF) over 3 years and a seasonal flooded forest (SFF) during 4 years using a light extinction method and two remote sensing methods (LAI MODIS product and the Landsat-METRIC method), with the objectives of (1) evaluating the performance of the remote sensing methods, and (2) understanding how total, overstory and understory LAI interact with micrometeorological variables. Total, overstory and understory LAI differed between both sites, with ASTF having higher LAI values than SFF, but neither site exhibited year-to-year variation in LAI despite large differences in meteorological variables. LAI values at the two sites have different patterns of correlation with micrometeorological variables. ASTF exhibited smaller seasonal variations in LAI than SFF. In contrast, SFF exhibited small changes in total LAI; however, dry season declines in overstory LAI were counteracted by understory increases in LAI. MODIS LAI correlated weakly to total LAI for SFF but not for ASTF, while METRIC LAI had no correlation to total LAI. However, MODIS LAI correlated strongly with overstory LAI for both sites, but had no correlation with understory LAI. Furthermore, LAI estimates based on canopy light extinction were correlated positively with seasonal variations in rainfall and soil water content and negatively with vapor pressure deficit and solar radiation; however, in some cases satellite-derived estimates of LAI exhibited no correlation with climate variables (METRIC LAI or MODIS LAI for ASTF). These data indicate that the satellite-derived estimates of LAI are insensitive to the understory variations in LAI that occur in many seasonal tropical forests and the micrometeorological variables that control seasonal variations in leaf phenology. While more ground-based measurements are needed to adequately quantify the performance of these satellite-based LAI products, our data indicate that their output must be interpreted with caution in seasonal tropical forests.     

Hydraulic acclimation to shading in boreal conifers of varying shade tolerance

The purpose of this study was to determine how shading affects the hydraulic and wood‐anatomical characteristics of four boreal conifers (Pinus banksiana, Pinus contorta, Picea glauca and Picea mariana) that differ in shade tolerance. Plants were grown in an open field and under a deciduous‐dominated overstory for 6 years. Sapwood‐ and leaf‐area specific conductivity, vulnerability curves, and anatomical measurements (light and scanning electron microscopy) were made on leading shoots from six to nine trees of each treatment combination. There was no difference in sapwood‐area specific conductivity between open‐grown and understory conifers, although two of four species had larger tracheid diameters in the open. Shaded conifers appeared to compensate for small diameter tracheids by changes in pit membrane structure. Scanning electron microscopy revealed that understory conifers had thinner margo strands, greater maximum pore size in the margo, and more torus extensions. All of these trends may contribute to inadequate sealing of the torus. This is supported by the fact that all species showed increased vulnerability to cavitation when grown in the understory. Although evaporative demand in an understory environment is low, a rapid change into fully exposed conditions could be detrimental for shaded conifers.     

热带次生林不同林层植物叶片非结构性碳水化合物的季节变化及其对氮磷添加的响应

非结构性碳水化合物(Non-structural Carbohydrates, NSCs)是植物生长代谢过程中重要的能量来源。通过在华南热带次生林进行氮磷添加试验,探究不同林层植物叶片NSCs的季节变化及其对氮磷添加的响应,取样时间为2019年1月、4月、7月和10月。结果表明:1)植物叶片NSCs存在显著的种间差异,磷(P)添加对叶片淀粉和NSCs含量具有显著影响,且物种与磷添加的交互作用显著影响叶片淀粉含量。2)黑嘴蒲桃和紫玉盘叶片NSCs含量对氮(N)添加的响应较为敏感,而白车和竹节叶片NSCs含量对P添加的响应较为敏感,氮磷同时添加(+NP)对植物叶片NSCs的增效作用最好。3)植物叶片NSCs存在显著的季节性变化,且季节与林层间的交互作用对叶片可溶性糖和NSCs含量具有显著影响。4)不同林层植物对氮磷添加的响应不同,氮磷添加使林下层植物叶片可溶性糖含量增高,林冠层降低,在干季,N添加会使林下层植物叶片淀粉含量增高,林冠层降低。P添加的影响恰好与之相反。在湿季,氮磷添加使林下层和林冠层植物叶片的淀粉含量增加。5)林冠层植物叶片NSCs含量高于林下层,且林下层植物叶片NSCs含量...     

Woodland canopy structure and the light response of juvenile Quercus lobata (Fagaceae)

I investigated competition for light between canopy plants and juvenile valley oaks (Quercus lobata Nee) in a mixed-broadleaf woodland of California's northern Coast Ranges. Canopy effects on understory light supply were separated among the overlying adult valley oak, the adult's woody understory, and neighboring trees and shrubs through a series of light sampling surveys and measurements of the number, size, and spatial distribution of neighboring plants. Light supply in the understory was primarily influenced by neighboring plants, with no detectable effect of the overlying adult valley oak. Light supply in the understory averaged 25% full sun due to a high frequency of canopy gaps and a typically open understory. Seedling response to understory light supply was investigated in an experimental sunfleck gradient (10%, 19%, and 100% full sun). Between 10% and 100% full sun, seedling growth increased by 90% and the shoot:∗∗∗root ratio changed from 1.561 to 0.607. Shade seedlings were also taller and produced fewer, larger, and thinner leaves than seedlings grown in full sun. A field survey of the spatial distribution and crown morphology of saplings and young adults found 1) the distance between young valley oaks and neighboring overstory trees to increase with neighbor size, and 2) crowns of the young oaks to be skewed away from neighbors. Although shading by the canopy was only moderate, canopy effects on understory light supply may restrict juvenile recruitment of valley oak in this woodland.     

天童国家森林公园常绿阔叶林不同演替阶段群落光环境特征比较

常绿阔叶林是我国中亚热带东部典型植被类型,根据野外踏查和固定样地调查发现,天童国家森林公园内发育着常绿阔叶林一个完整的演替系列,包含着6个不同演替阶段。应用W inScanopy For C anopy A na lys is软件对研究区内不同演替阶段群落冠层进行分析,得到不同群落冠层和林下的光环境特征指标:PPFD(光合光量子通量密度)和相关的冠层结构形态学指标G ap fraction(空隙度)、LA I(叶面积指数)、M LA(平均叶倾角),通过对这些指标的分析比较,得到的基本规律大致是林冠层的光合有效光量子通量密度随演替逐渐降低,林冠下面的光合有效光量子通量密度随着群落演替的进展变化更为明显。马尾松林的林冠空隙度明显高于其他阶段的群落,总的趋势是随群落演替的进展而降低。叶面积指数随演替的进展而呈增加趋势。平均叶倾角随演替的进展先增大而后减小。这些结果反映了常绿阔叶林不同演替阶段群落由于不同树种树冠形态学结构的差异和微环境的不同,形成了特定群落内的特定光环境。