Response of macroalgal assemblages from rockpools to climate change: effects of persistent increase in temperature and CO2

Anthropogenically induced global climate change has important implications for marine ecosystems with unprecedented ecological and economic consequences. Climate change will include the simultaneous increase of temperature and CO₂ concentration in oceans. However, experimental manipulations of these factors at the community scale are rare. In this study, we used an experimental approach in mesocosms to analyse the combined effects of elevated CO₂ and temperature on macroalgal assemblages from intertidal rock pools. Our model systems were synthetic assemblages of varying diversity and understory component and canopy species identity. We used assemblages invaded by the non‐indigenous canopy forming alga Sargassum muticum and assemblages with the native canopy species Cystoseira tamariscifolia. We examined the effects of both climate change factors on several ecosystem functioning variables (i.e. photosynthetic efficiency, productivity, respiration and biomass) and how these effects could be shaped by the diversity and species identity of assemblages. CO₂ alone or in combination with temperature affected the performance of macroalgae at both individual and assemblage level. In particular, high CO₂ and high temperature (20°C) drastically reduced the biomass of macroalgal assemblages and affected their productivity and respiration rates. The identity of canopy species also played an important role in shaping assemblage responses, whereas species richness did not seem to affect such responses. Species belonging to the same functional effect group responded differently to the same environmental conditions. Data suggested that assemblages invaded with S. muticum might be more resistant in a future scenario of climate change. Thus, in a future scenario of increasing temperature and CO₂ concentration, macroalgal assemblages invaded with canopy‐forming species sharing response traits similar to those of S. muticum could be favoured.     

Importance of the understory stratum to entomofaunal diversity in a temperate deciduous forest

The vertical stratification of lepidopteran and coleopteran communities in a cool-temperate deciduous forest in Japan was examined to evaluate the hypothesis of an expected uniform distribution of mobile flying insects between the canopy and understory of temperate forests. Lepidopteran and coleopteran insects were trapped using light traps at three sites in each of the canopy and understory for three consecutive nights each month from April to October 2001. For Lepidoptera, species richness, abundance, and family richness were significantly higher in the understory than in the canopy. For Coleoptera, only abundance was larger in the canopy relative to the understory; species and family richness did not differ between the strata. The beta diversity of the lepidopteran community was larger between the strata than among sites, but the coleopteran community showed an inverse pattern. These results imply the presence of vertical stratification within the lepidopteran community, but not within the coleopteran community, in the temperate forest. The understory contributes more than the canopy to lepidopteran diversity in the temperate forest, although this stratification may be relatively weak because, in contrast to the situation in tropical forests, the canopy and understory assemblages share many species.     

How Does Restoration of Native Canopy Affect Understory Vegetation Composition? Evidence from Riparian Communities of the Hunter Valley Australia

Restoration of native vegetation often focuses on the canopy layer species, with the assumption that regeneration of the understory elements will occur as a consequence. The goal of this study was to assess the influence of canopy restoration on the composition and abundance of understory plant species assemblages along riparian margins in the Hunter Valley, NSW, Australia. We compared the floristic composition (richness, abundance, and diversity) of understory species between nonrevegetated (open) and canopy revegetated plots across five sites. A number of other factors that may also influence understory vegetation, including soil nutrients, proximity to main channel, and light availability, were also measured. We found that sites where the canopy had been restored had lower exotic species richness and abundance, as well as higher native species cover, but not native species richness, compared with open sites. Multivariate analysis of plots based on plant community composition showed that revegetated sites were associated with lower total species diversity, light availability, and exotic cover. This study has found that the restoration of the canopy layer does result in lower exotic species richness and cover, and higher native species cover and diversity in the understory, a desirable restoration outcome. Our results provide evidence that restoration of native canopy species may facilitate restoration of native understory species; however, other interventions to increase native species richness of the understory should also be considered as part of management practice.     

伏牛山自然保护区森林冠层结构对林下植被特征的影响

在伏牛山自然保护区典型地段设立样方,测定了森林生态系统内几种典型群落类型的冠层结构、光环境特征,调查了林下植被的特征,分析了它们之间的相互关系.结果显示:各群落的冠层结构和光环境有一定的差异,单因素方差分析表明,部分群落间的差异性达到显著水平;各群落灌木层物种丰富度、多样性和均匀度均高于草本层,而优势度正相反;线性拟合的结果表明,草本层的物种丰富度、多样性与冠下光合量子通量密度间呈极显著负相关,优势度与冠下光合量子通量密度间呈显著正相关,灌木层各参数与冠层结构特征间相关性不显著.研究表明,冠层结构的变化对草本层(包括更新幼苗)的影响显著高于灌木层.林隙/林窗或林中空地的出现可能对草本物种或其他阳性及先锋物种具有促进作用,而对优势种幼苗的萌发和定植产生负效应.推测在典型的落叶阔叶林生态系统演替进程中,林下光照强度可能不是最主要的限制因素,优势种种子的扩散、萌发和定植限制可能更重要.     

Patterns and controls of the dynamics of net primary production by understory macroalgal assemblages in giant kelp forests

Macroalgae are important primary producers in many subtidal habitats, yet little information exists on the temporal and spatial dynamics of net primary production (NPP) by entire subtidal assemblages. This knowledge gap reflects the logistical challenges in measuring NPP of diverse macroalgal assemblages in shallow marine habitats. Here, we couple a simple primary production model with nondestructive estimates of taxon‐specific biomass on subtidal reefs off Santa Barbara, California to produce a 4‐year time series of net primary production by intact assemblages of understory macroalgae in giant kelp forests off Santa Barbara, California, USA. Daily bottom irradiance varied significantly throughout the year, and algal assemblages were on average exposed to saturating irradiance for only 1.3–4.5 h per day, depending on the time of year. Despite these variable light‐limiting conditions, biomass rather than irradiance explained the vast majority of variation observed in daily NPP at all times of the year. Measurements of peak biomass in spring and summer proved to be good predictors of NPP for the entire year, explaining as much as 76% of the observed variation. In contrast, bottom irradiance was a poor predictor of NPP, explaining <10% of the variation in NPP when analyzed seasonally and ~2% when evaluated annually. Our finding that annual NPP by macroalgal assemblages can be predicted from a single, nondestructive measurement of biomass should prove useful for developing time series data that are necessary to evaluate natural and anthropogenic changes in NPP by one of the world's most productive ecosystems.     

Crown architecture in sun and shade environments: assessing function and trade-offs with a three-dimensional simulation model

Sun and shade environments place markedly different constraints on the photosynthetic performance of plants. Leaf-level photosynthetic responses to sun and shade have been extensively investigated, whereas there has been much less research on the functional role of crown architecture in these environments. This paper focuses on the role of architecture in maximizing light capture and photosynthesis in shaded understories and in minimizing exposure to excess radiation in open high light environments. Understanding these contrasting roles of architecture is facilitated by application of a three-dimensional structural-functional model, Y-plant. Surveys of understory plants reveal a diversity of architectures but a strong convergence at only modest light-capture efficiencies because of significant self-shading. Simulations with Psychotria species revealed that increasing internode lengths would increase light-capture efficiencies and whole plant carbon gain. However, the costs of the additional required biomechanical support was high, which, in terms of relative growth rates, would override the advantage provided by higher light-capture efficiencies. In high light environments, leaf angles and self-shading provide structural photoprotection, minimizing potential damage from photoinhbition. Simulations reveal that without these structural protections photoinhibition of photosynthesis is likely to be much greater with daily carbon gain significantly reduced.     

Canopy photosynthesis of six major arable crops is enhanced under diffuse light due to canopy architecture

Diffuse radiation generally increases photosynthetic rates if total radiation is kept constant. Different hypotheses have been proposed to explain this enhancement of photosynthesis, but conclusive results over a wide range of diffuse conditions or about the effect of canopy architecture are lacking. Here, we show the response of canopy photosynthesis to different fractions of diffuse light conditions for five major arable crops (pea, potato, wheat, barley, rapeseed) and cover crops characterized by different canopy architecture. We used 13 years of flux and microclimate measurements over a field with a typical 4 year crop rotation scheme in Switzerland. We investigated the effect of diffuse light on photosynthesis over a gradient of diffuse light fractions ranging from 100% diffuse (overcast sky) to 11% diffuse light (clear‐sky conditions). Gross primary productivity (GPP) increased with diffuse fraction and thus was greater under diffuse than direct light conditions if the absolute photon flux density per unit surface area was kept constant. Mean leaf tilt angle (MTA) and canopy height were found to be the best predictors of the diffuse versus direct radiation effect on photosynthesis. Climatic factors, such as the drought index and growing degree days (GDD), had a significant influence on initial quantum yield under direct but not diffuse light conditions, which depended primarily on MTA. The maximum photosynthetic rate at 2,000 µmol m^?2 s^?1 photosynthetically active radiation under direct conditions strongly depended on GDD, MTA, leaf area index (LAI) and the interaction between MTA and LAI, while under diffuse conditions, this parameter depended mostly on MTA and only to a minor extent on canopy height and their interaction. The strongest photosynthesis enhancement under diffuse light was found for wheat, barley and rapeseed, whereas the lowest was for pea. Thus, we suggest that measuring canopy architecture and diffuse radiation will greatly improve GPP estimates of global cropping systems.     

Metabolic compensation constrains the temperature dependence of gross primary production

Gross primary production (GPP) is the largest flux in the carbon cycle, yet its response to global warming is highly uncertain. The temperature dependence of GPP is directly linked to photosynthetic physiology, but the response of GPP to warming over longer timescales could also be shaped by ecological and evolutionary processes that drive variation in community structure and functional trait distributions. Here, we show that selection on photosynthetic traits within and across taxa dampens the effects of temperature on GPP across a catchment of geothermally heated streams. Autotrophs from cold streams had higher photosynthetic rates and after accounting for differences in biomass among sites, biomass‐specific GPP was independent of temperature in spite of a 20 °C thermal gradient. Our results suggest that temperature compensation of photosynthetic rates constrains the long‐term temperature dependence of GPP, and highlights the importance of considering physiological, ecological and evolutionary mechanisms when predicting how ecosystem‐level processes respond to warming.     

Understory Plant Community Composition Is Associated with Fine-Scale Above- and Below-Ground Resource Heterogeneity in Mature Lodgepole Pine (Pinus contorta) Forests

Understory plant communities play critical ecological roles in forest ecosystems. Both above- and below-ground ecosystem properties and processes influence these communities but relatively little is known about such effects at fine (i.e., one to several meters within-stand) scales, particularly for forests in which the canopy is dominated by a single species. An improved understanding of these effects is critical for understanding how understory biodiversity is regulated in such forests and for anticipating impacts of changing disturbance regimes. Our primary objective was to examine the patterns of fine-scale variation in understory plant communities and their relationships to above- and below-ground resource and environmental heterogeneity within mature lodgepole pine forests. We assessed composition and diversity of understory vegetation in relation to heterogeneity of both the above-ground (canopy tree density, canopy and tall shrub basal area and cover, downed wood biomass, litter cover) and below-ground (soil nutrient availability, decomposition, forest floor thickness, pH, and phospholipid fatty acids (PLFAs) and multiple carbon-source substrate-induced respiration (MSIR) of the forest floor microbial community) environment. There was notable variation in fine-scale plant community composition; cluster and indicator species analyses of the 24 most commonly occurring understory species distinguished four assemblages, one for which a pioneer forb species had the highest cover levels, and three others that were characterized by different bryophyte species having the highest cover. Constrained ordination (distance-based redundancy analysis) showed that two above-ground (mean tree diameter, litter cover) and eight below-ground (forest floor pH, plant available boron, microbial community composition and function as indicated by MSIR and PLFAs) properties were associated with variation in understory plant community composition. These results provide novel insights into the important ecological associations between understory plant community composition and heterogeneity in ecosystem properties and processes within forests dominated by a single canopy species.     

Importance of canopy structure on photosynthesis in single- and multi-species assemblages of marine macroalgae

Plant communities utilize available irradiance with different efficiency depending not only on their photosynthetic characteristics but also on the canopy structure and density. The importance of canopy structure are well studied in terrestrial plant communities but poorly studied in aquatic plant communities. The objective of this study was to evaluate macroalgal community photosynthesis in artificial constructed communities of one to four species with different morphologies along a range of leaf (i.e.=thallus) area densities. In a laboratory set-up we measured net photosynthesis and dark respiration in constructed assemblages of macroalgae, excluding effects other than photosynthesis of individual tissue and distribution of photons in the canopy from influencing metabolism. We hypothezised that 1) canopy structure determines the actual rates of photosynthesis relative to the optimal rates and 2) multi-species communities attain higher maximum photosynthetic rates than single species communities. We found that differences in canopy structure outweighed large differences in tissue photosynthesis resulting in relatively similar maximum community photosynthetic rates among the different single and multi-species assemblages (20.1–40.5 μmol O₂ m⁻² s⁻¹). Canopy structure influenced community photosynthesis both at low and high leaf area densities because it determines the ability of macroalgae to use the photosynthetic potential of their individual tissues. Due to an averaging effect the photosynthetic rate at high leaf area density was more similar among multi-species community than among single-species communities. Multi-species communities had, on average, a slightly higher photosynthetic production than expected from photosynthesis of single species communities. Moreover multi-species communities were capable of exposing new tissue to irradiance up to high densities thereby avoiding a decrease in net photosynthesis. This finding suggests that multi-species communities may be able to maintain higher biomass per unit ground area than single-species communities.     

Impact of nitrogen allocation on growth and photosynthesis of Miscanthus (Miscanthus × giganteus)

Nitrogen (N) addition typically increases overall plant growth, but the nature of this response depends upon patterns of plant nitrogen allocation that vary throughout the growing season and depend upon canopy position. In this study seasonal variations in leaf traits were investigated across a canopy profile in Miscanthus (Miscanthus × giganteus) under two N treatments (0 and 224 kg ha^?1) to determine whether the growth response of Miscanthus to N fertilization was related to the response of photosynthetic capacity and nitrogen allocation. Miscanthus yielded 24.1 Mg ha^?1 in fertilized plots, a 40% increase compared to control plots. Photosynthetic properties, such as net photosynthesis (A), maximum rate of rubisco carboxylation (V_cmax), stomatal conductance (g_s) and PSII efficiency (F_v'/F_m'), all decreased significantly from the top of the canopy to the bottom, but were not affected by N fertilization. N fertilization increased specific leaf area (SLA) and leaf area index (LAI). Leaf N concentration in different canopy layers was increased by N fertilization and the distribution of N concentration within canopy followed irradiance gradients. These results show that the positive effect of N fertilization on the yield of Miscanthus was unrelated to changes in photosynthetic rates but was achieved mainly by increased canopy leaf area. Vertical measurements through the canopy demonstrated that Miscanthus adapted to the light environment by adjusting leaf morphological and biochemical properties independent of nitrogen treatments. GPP estimated using big leaf and multilayer models varied considerably, suggesting a multilayer model in which V_cmax changes both through time and canopy layer could be adopted into agricultural models to more accurately predict biomass production in biomass crop ecosystems.     

Survival and performance of the invasive vine Vincetoxicum rossicum (Apocynaceae) from seeds of different embryo number under two light environments

The nonnative vine Vincetoxicum rossicum threatens several ecosystems in the Lower Great Lakes Basin of North America. One feature that may contribute to its invasiveness is the production of some seeds with multiple embryos (polyembryony), which may be beneficial as a bet-hedging strategy in variable environments. However, lower seed reserves per embryo in polyembryonic seeds may entail costs in low-light environments. The effect of seed from three embryonic classes (1, 2, or 3 embryos/seed) on V. rossicum survival and growth was studied under two forest understory light environments: full canopy (shade) or canopy gaps (light) in New York state. Two seedling cohorts were planted, in May 2004 and in May 2005. The survival and growth of seedlings was monitored biweekly for two (2005 cohort) or three (2004 cohort) seasons. For both cohorts, plants grown in canopy shade had reduced survival and growth compared with those grown in gaps. Contrary to expectations, seed embryo number had no effect on the final height, survival, or dry mass of plants in either habitat. Our results suggest that any fitness advantage provided by polyembryony may be habitat (light) dependent and not a general trait that affords V. rossicum a benefit in all habitats colonized.     

西双版纳热带雨林林窗空间分布格局及其特征数与林窗下植物多样性的相关性

林窗作为森林群落中一种重要的干扰方式, 对林下物种构成有着重要的影响。开展林窗空间格局及其特征指数与林下植物多样性关系研究对于探讨林窗对林下生物多样性的影响有重要意义, 有助于进一步了解群落动态, 在物种多样性保护方面也具有指导作用。本研究在西双版纳热带雨林地区随机选取3块大小为1 ha的热带雨林为研究样地, 采用轻小型六旋翼无人机搭载Sony ILCE-A7r可见光传感器, 分别获取各个样地的高清数字影像, 结合数字表面高程模型以及各个样地的地形数据用以确定各样区的林窗分布格局, 并进一步提取出各林窗的景观格局指数。结合地面样方基础调查数据, 对各样地各林窗下植物多样性情况进行统计, 旨在分析热带雨林林窗空间分布格局以及林窗下植物多样性对各林窗空间格局特征的响应情况。研究表明, 西双版纳州热带雨林林窗呈大而分散的空间分布, 林窗空间格局特征指数如林窗形状复杂性指数、林窗面积都与林下植物多样性呈显著正相关关系。在面积小的林窗下, 较之林窗形状复杂性因子, 林窗面积大小对林下植物多样性影响更显著; 在面积达到一定程度后, 相对于面积因子, 林窗形状复杂性指数对林下植物多样性影响更显著, 各样地林窗皆趋于向各自所处样地顶极群落发展。     

遮荫处理对梅叶冬青叶片形态、光合特性和生长的影响

为了解遮荫环境对梅叶冬青(Ilex asprella)生长和光合特性的影响,采用遮荫网的方法模拟85%、56%和全光照等3种光照环境,研究了遮荫对其生长、光合参数以及生物量等的影响。结果表明,经一年遮荫处理后,梅叶冬青的叶绿素a、b和叶绿素总量都随着遮荫强度的增加而显著增高,胡萝卜素含量则显著降低。与对照相比,56%遮荫处理显著提高梅叶冬青的最大净光合速率和光饱和点,分别提高了17.6%和25.2%,但是85%遮阴处理则显著降低最大净光合速率和光饱和点,分别降低了18.2%和24.1%,两种遮荫处理均显著降低了光补偿点。叶长、叶宽、比叶面积、单片叶面积和叶片含水量均随着遮荫强度的增加而显著增加,而叶片厚度则显著减小。遮荫处理明显抑制整株生物量增长,减小根冠比,但是株高、冠幅和径向生长随遮荫处理时间不同而有所变异。因此,梅叶冬青有耐荫偏阳的特性,在林下种植时需及时调控乔冠层的透光率,一般应大于44%。     

Photosynthetic responses of Miconia species to canopy openings in a lowland tropical rainforest

We examined the photosynthetic acclimation of three tropical species of Miconia to canopy openings in a Costa Rican rainforest. The response of photosynthesis to canopy opening was very similar in Miconia affinis, M. gracilis, and M. nervosa, despite differences in growth form (trees and shrubs) and local distributions of plants (understory and gap). Four months after the canopy was opened by a treefall, photosynthetic capacity in all three species had approximately doubled from closed canopy levels. There were no obvious signs of high light damage after treefall but acclimation to the gap environment was not immediate. Two weeks after treefall, A_max, stomatal conductance, apprarent quantum efficiency, and dark respiration rates had not changed significantly from understory values. The production of new leaves appears to be an important component of light acclimation in these species. The only variables to differ significantly among species were stomatal conductance at A_max and the light level at which assimilation was saturated. M. affinis had a higher stomatal conductance which may reduce its water use efficiency in gap environments. Photosynthesis in the more shade-tolerant M. gracilis saturated at lower light levels than in the other two species. Individual plant light environments were assessed after treefall with canopy photography but they explained only a small fraction of plant variation in most measures of photosynthesis and growth. In conclusion, we speculate that species differences in local distribution and in light requirements for reproduction may be more strongly related to species differences in carbon allocation than in carbon assimilation.     

Photosynthetic acclimation of the liana Stigmaphyllon lindenianum to light changes in a tropical dry forest canopy

Tropical plant canopies show abrupt changes in light conditions across small differences in spatial and temporal scales. Given the canopy light heterogeneity, plants in this stratum should express a high degree of plasticity, both in space (allocation to plant modules as a function of opportunity for resource access) and time (photosynthetic adjustment to temporal changes in the local environment). Using a construction crane for canopy access, we studied light acclimation of the liana Stigmaphyllon lindenianum to sun and shade environments in a tropical dry forest in Panama during the wet season. Measured branches were randomly distributed in one of four light sequences: high- to low-light branches started the experiment under sun and were transferred to shade during the second part of the experiment; low- to high-light branches (LH) were exposed to the opposite sequence of light treatments; and high-light and low-light controls , which were exposed only to sun and shade environments, respectively, throughout the experiment. Shade branches were set inside enclosures wrapped in 63% greenhouse shade cloth. After 2 months, we transferred experimental branches to opposite light conditions by relocating the enclosures. Leaf mortality was considerably higher under shade, both before and after the transfer. LH branches reversed the pattern of mortality by increasing new leaf production after the transfer. Rates of photosynthesis at light saturation, light compensation points, and dark respiration rates of transferred branches matched those of controls for the new light treatment, indicating rapid photochemical acclimation. The post-expansion acclimation of sun and shade foliage occurred with little modification of leaf structure. High photosynthetic plasticity was reflected in an almost immediate ability to respond to significant changes in light. This response did not depend on the initial light environment, but was determined by exposure to new light conditions. Stigmaphyllon responded rapidly to light changes through the functional adjustment of already expanded foliage and an increase in leaf production in places with high opportunity for carbon gain. Received: 24 April 1998 / Accepted: 11 May 1999     

不同透光环境下红松光合色素含量的季节变动及应对策略

研究了辽东山区天然次生林内3种不同透光环境(强度透光、中度透光和弱度透光)下红松针叶光合色素(叶绿素a(Chl a)、叶绿素b(Chl b)、类胡萝卜素(Car)和叶绿素总量(Chl T))应对光环境季节变动做出的适应性调整。结果表明,随季节的变动(从春季至秋季),林分透光孔隙度逐渐减小。春季,透光度越大,红松叶绿素含量越高,Chl a/b值升高,Car/Chl T值降低;夏季,不同透光条件对红松光合色素含量无影响;秋季,各类透光条件下红松光合色素含量总体表现为升高的趋势,强度透光与中度透光条件红松针叶Chl a/b显著大于弱度透光,3种透光条件下红松Car/Chl T均降低。在春季红松开始生长前进行适当抚育,能提高光合色素含量,增强光合作用能力,促进生长。     

杉木人工林林下植物多样性及其环境对不同间伐强度的短期响应

以江西省官山林场的杉木人工林为研究对象,探究不同间伐强度（0%、20%、40%）物种多样性、地上生物量、林分光环境和土壤理化性质及其之间的相关性。结果表明：（1）林下物种数量随间伐强度的增加而增加,林下灌木层优势种不断变化,而草本层一直不变。（2）林下植物各多样性指数均随间伐强度的增大而增大,除Margalef指数外,其余各指数均在间伐40%与未间伐间呈显著差异。（3）灌木层地上生物量占总生物量的主体,且随间伐强度的增大而增大,不同间伐强度间差异显著,而草本层却呈相反趋势。（4）叶面积指数随间伐强度的增加而下降,冠层开度、林下直射光、散射光、总光照随之上升,但仅在间伐40%后显著。（5）土壤全氮含量间伐后显著上升,但土壤磷、钾、有机质含量均显著下降（6）灌木层多样性指数与灌木层地上生物量、土壤全氮呈显著正相关,与草本层地上生物量、速效磷、全钾含量呈显著负相关。草本层多样性指数与冠层开度、林下直射光、林下散射光、林下总光照呈显著正相关,与草本层地上生物量、叶面积指数呈显著负相关。总之,林下植物地上生物量、土壤化学性质是影响灌木层多样性指数变化的主要控制因子,林分光环境是影响草本层的主要控制因子。就本研究3种间伐强度而言,该地杉木人工林的最适宜间伐强度为40%。     

Ecophysiological responses of Fagus sylvatica seedlings to changing light conditions. II. The interaction of light environment and soil fertility on seedling physiology

The survival and growth of natural beech regeneration after canopy removal is variable and little is known about ecophysiological mechanisms of these responses. Biomass, nonstructural carbohydrate levels and nitrogen concentrations were measured in an Italian population of European beech seedlings. Seedlings were container-grown in two types of soil, organic and mineral, collected at the study site. The seedlings were grown under three light treatments: under full beech canopy (understory), exposed to full sun only during midday (gap) and under full sun (clearing). Leaf gas exchange and chlorophyll a fluorescence parameters were measured and then foliar analyses were conducted for chlorophyll, phenolic and tannin levels. Biomass and allocation were significantly affected by light and soil treatments. The clearing seedlings and those in organic soil were larger than seedlings in the other light treatments or soil type. Total nonstructural carbohydrate concentrations were lower in the understory seedlings and significant differences between soil types were present in the gap and clearing seedlings. Nitrogen concentrations were higher in the understory seedlings and those growing in the organic soil compared to the other treatments. Gas exchange rates were highest in clearing and the organic soil seedlings. Gap seedlings exhibited photosynthetic acclimation that allowed them to utilize high light of midday and any sunflecks during the morning and afternoon. Relative fluorescence was significantly influenced by both light treatment and soil type, with the highest values observed in the gap seedlings. Light response curves showed decreasing apparent maximum quantum efficiency from the understory to clearing, while maximum photosynthetic rate was highest in the gap seedlings. Chlorophyll concentration was highest in understory seedlings and those growing in organic soil and higher in seedlings growing in organic than in mineral soil. Both foliar tannin and phenolic levels were highest in clearing seedlings, and only tannin concentrations were affected by soil type. Understory seedlings had the highest mortality and insect herbivory; the latter was found to be inversely related to tannin concentration. Overall, growth and photosynthesis in beech seedlings responded positively to high light associated with small canopy gaps. Organic soil increased seedling size, particularly in the gap and clearing environments. We conclude that forest gaps are favorable for photosynthesis and growth of European beech seedlings.     

Linking light attenuation,sunflecks, and canopy architecture in mesic shrub thickets

Expansion of shrubs into grasslands is often accompanied by a reduction in understory light and an associated reduction of shade-intolerant species. However, effects of specific canopy architectural characteristics on the light environment under shrub thickets are unknown. Our objective was to determine what characteristics of canopy architecture most influence understory light in monospecific shrub thickets. We quantified understory light and canopy architecture for five shrub species in the eastern United States that have a history of expansion, and we used multiple regression to determine which canopy characteristics best predicted light attenuation and relative contribution of sunflecks. Measurements included leaf angle, leaf azimuth, branch bifurcation ratio, leaf area index (LAI), canopy depth (the vertical distance from the bottommost leaf to the top of the canopy), and leaf area density (LAD) as well as understory photosynthetic photon flux density (PPFD). The best predictor of light attenuation and the occurrence of sunflecks for all species was canopy depth. Multiple leaf and plant-level traits were correlated with canopy depth but not with LAI or LAD. The invasive shrub Elaeagnus umbellata had the lowest understory light levels of the species examined although LAI values for Morella cerifera and Rhododendron maximum were higher. Branch bifurcation ratios for E. umbellata were significantly higher than for other species and this likely contributed to the differences in light attenuation and suppression of sunflecks. The potential of shrubs to intercept light is primarily dependent on vertical distribution of leaves in the canopy which is itself correlated with fine-scale, species-specific variations in leaf display.