The compensatory responses of an understory herb to experimental damage are habitat-dependent

Canopy gap formation strongly influences the diversity and dynamics of both tropical and temperate forests. It is often viewed as inherently beneficial for understory plants, primarily because growth and flowering are enhanced when light is no longer a limiting resource. It can also be detrimental, however, because plants can be damaged by falling crowns or branches. To elucidate the responses of the Amazonian understory herb Heliconia acuminata to damage sustained during gap formation, we transplanted both experimentally damaged and control plants to canopy gaps and the forest understory. We then measured their patterns of growth and biomass allocation 10 mo later. Despite losing approximately 50% of their leaf area, all damaged plants survived the duration of our experiment. Furthermore, damaged plants transplanted to gaps had relative growth rates that far exceeded those of undamaged plants in both gaps and the forest understory. There were also significant interactions between damage and destination habitat type on root to shoot ratios and leaf-area ratios. Our results suggest the ability of herbaceous plants to recover from damage, as well as patterns of post-damage biomass allocation, may be habitat-dependent in ways that have previously remained unexplored.     

Response of clonal plasticity of Fargesia nitida to different canopy conditions of subalpine coniferous forest

The aim of this study is to explore the effects of canopy conditions on clump and culm numbers, and the morphological plasticity and biomass distribution patterns of the dwarf bamboo species Fargesia nitida. Specifically, we investigated the effects of canopy condi-tions on the growth and morphological characteristics of F. nitida, and the adaptive responses of F. nitida to dif-ferent canopy conditions and its ecological senses. The results indicate that forest canopy had a significant effect on the genet density and culm number per clump, while it did not affect the ramet density. Clumps tended to be few and large in gaps and forest edge plots, and small under forest understory plots. The ramets showed an even distribution under the closed canopy, and clus-ter distribution under gaps and forest edge plots. The forest canopy had a significant effect on both the ramets'biomass and biomass allocation. Favourable light conditions promoted ramet growth and biomass accumulation. Greater amounts of biomass in gaps and forest edge plots were shown by the higher number of culms per clump and the diameter of these culms. Under closed canopy, the bamboos increased their branching angle, leaf biomass allocation, specific leaf area and leaf area ratio to exploit more favourable light conditions in these locations. The spacer length, specific spacer length and spacer branching angles all showed significant differences between gaps and closed canopy conditions. The larger specific spacer length and spacer branching angle were beneficial for bamboo growth, scattering the ramets and exploiting more favourable light conditions. In summary, this study shows that to varying degrees, F nitida exhibits both a wide ecological amplitude and high degree of morphological plasticity in response to differing forest canopy conditions. More-over, the changes in plasticity enable the plants to optimize their light usage efficiency to promote growth and increase access to resources available in heteerogeneous light environoments.     

Response of clonal plasticity of Fargesia nitida to different canopy conditions of subalpine coniferous forest

The aim of this study is to explore the effects of canopy conditions on clump and culm numbers, and the morphological plasticity and biomass distribution patterns of the dwarf bamboo species Fargesia nitida. Specifically, we investigated the effects of canopy conditions on the growth and morphological characteristics of F. nitida, and the adaptive responses of F. nitida to different canopy conditions and its ecological senses. The results indicate that forest canopy had a significant effect on the genet density and culm number per clump, while it did not affect the ramet density. Clumps tended to be few and large in gaps and forest edge plots, and small under forest understory plots. The ramets showed an even distribution under the closed canopy, and cluster distribution under gaps and forest edge plots. The forest canopy had a significant effect on both the ramets’ biomass and biomass allocation. Favourable light conditions promoted ramet growth and biomass accumulation. Greater amounts of biomass in gaps and forest edge plots were shown by the higher number of culms per clump and the diameter of these culms. Under closed canopy, the bamboos increased their branching angle, leaf biomass allocation, specific leaf area and leaf area ratio to exploit more favourable light conditions in these locations. The spacer length, specific spacer length and spacer branching angles all showed significant differences between gaps and closed canopy conditions. The larger specific spacer length and spacer branching angle were beneficial for bamboo growth, scattering the ramets and exploiting more favourable light conditions. In summary, this study shows that to varying degrees, F. nitida exhibits both a wide ecological amplitude and high degree of morphological plasticity in response to differing forest canopy conditions. Moreover, the changes in plasticity enable the plants to optimize their light usage efficiency to promote growth and increase access to resources available in heterogeneous light environments. __________ Translated from Acta Ecologica Sinica, 2006, 26(12): 4019–4026 [译自: 生态学报]     

Light gradient partitioning by tropical tree seedlings in the absence of canopy gaps

To explore the importance of light availability for seedling growth in low light environments, we examined light-dependent growth, biomass allocation and mortality of tree seedlings growing in sites with 0.2-6.5% full sun, the range of light commonly encountered in the understory of closed canopy, lowland tropical forests. We transplanted seedlings of the canopy tree species, Dipteryx panamensis, Virola koschnyii, and Brosimum alicastrum into second-growth forest and native tree plantations at La Selva Biological Station, Costa Rica. We assessed seedling survival, growth, and seedling light environments bimonthly for 14 months. Plants were harvested at the end of the study to assess leaf area, total biomass, biomass partitioning and root growth. Survivorship of all species exceeded 60% across all microsites, although both D. panamensis and B. alicastrum had lower probabilities of survival in the darkest microsites. All species showed a strong positive relationship between light availability and growth, increasing in total biomass as light increased. However, the strength of the growth response differed among species causing a change in the rank order of species growth rates as light availability increased. Although D. panamensis showed the lowest growth rates in the darkest microsites, a strong response to increasing light led to a cross-over in performance, such that D. panamensis had the highest growth rate at the highest light levels studied. These data suggest that resource gradient partitioning could occur even in low light environments (0.2-6.5%). Given the limited range of light regimes sampled (i.e., non-gap microsites), our data demonstrate that growth of tropical tree seedlings beneath closed canopies is highly sensitive to light availability and that shade-tolerant species vary in these responses. Our results show that understory light heterogeneity, in the absence of canopy gaps, can significantly affect recruitment processes for shade-tolerant tree species.     

亚高山暗针叶林不同林冠环境下华西箭竹的克隆可塑性

以亚高山暗针叶林3种林冠环境中以及暗针叶林林缘的华西箭竹（Fargesia nitida）为对象,对其无性系数量特征、无性系根茎特征、分株生物量以及分株形态特征进行了对比研究。结果表明：（1）林冠环境的差异导致了不同种群的基株密度和每基株分株数的显著差异,但林冠环境差异不影响分株密度。林冠郁闭度愈大,每基株分株数愈少,分株分布愈均匀。（2）不同林冠环境间。分株生物量、分株构件生物量和分株构件的生物量分配百分率均有显著差异。开敞的林冠环境有利于华西箭竹的生长和生物量积累。（3）随着林冠郁闭度的增加,华西箭竹通过增大分枝角度、叶生物量分配百分率、比叶面积和叶面积率以提高光能利用效率,有效适应弱光环境。（4）隔离者长度、隔离者直径和分枝强度在林缘和林窗环境中要显著大于林内环境;同级隔离者分枝角度随林冠郁闭度的增加而最大,其值在林下显著大于林窗和林缘,而异级隔离者分枝角度的变化则正好相反。研究表明,华西箭竹种群在不同的林冠环境中发生了明显的可塑性变化,这些可塑性变化是种群对林冠郁闭度差异的适应性反应的结果,有利于增强种群对环境中有限光资源的利用。     

根茎型植物扁秆荆三棱对光照强度和养分水平的生长响应及资源分配策略

自然界中光照和养分因子常存在时空变化,对植物造成选择压力。克隆植物可通过克隆生长和生物量分配的可塑性来适应环境变化。尽管一些研究关注了克隆植物对光照和养分因子的生长响应,但尚未深入全面了解克隆植物对光照和养分资源投资的分配策略。以根茎型草本克隆植物扁秆荆三棱(Bolboschoenus planiculmis)为研究对象,在温室实验中,将其独立分株种植于由2种光照强度(光照和遮阴)和4种养分水平(对照、低养分、中养分和高养分)交叉组成的8种处理组合中,研究了光照和养分对其生长繁殖及资源贮存策略的影响。结果表明,扁秆荆三棱的生长、无性繁殖及资源贮存性状均受到光照强度的显著影响,在遮阴条件下各生长繁殖性状指标被抑制。且构件的数目、长度等特征对养分差异的可塑性响应先于其生物量积累特征。在光照条件下,高养分处理的总生物量、叶片数、总根茎分株数、长根茎分株数、总根茎长、芽长度、芽数量等指标大于其他养分处理,而在遮阴条件下,其在不同养分处理间无显著差异,表明光照条件可影响养分对扁秆荆三棱可塑性的作用,且高营养水平不能补偿由于光照不足而导致的生长能力下降。光照强度显著影响了总根茎、总球茎及大、中、小球茎的生物量分配,遮阴条件下,总生物量减少了对地下部分根茎和球茎的分配,并将有限的生物量优先分配给小球茎。总根茎的生物量分配未对养分发生可塑性反应,而随着养分增加,总球茎分配下降,说明在养分受限的环境中球茎的贮存功能可缓冲资源缺乏对植物生长的影响。在相同条件下,根茎生物量对长根茎的分配显著大于短根茎,以保持较高的繁殖能力;而总球茎对有分株球茎的生物量分配小于无分株球茎,表明扁秆荆三棱总球茎对贮存功能的分配优先于繁殖功能。研究为进一步理解根茎型克隆植物对光强及基质养分环境变化的生态适应提供了依据。     

Phenotypic plasticity of lianas in response to altered light environment

The growth, morphology and biomass allocation of 11 liana species (six light-demanding and five shade-tolerant) were investigated by growing plants in three contrasting light environments (i.e., field, forest edge and forest interior). Our objectives were to determine: (1) changes in plant traits at the species level; and (2) differences in light-demanding and shade-tolerant species in response to altered light environment. We found that all seedlings of liana species increased in total biomass, total leaf area, relative growth rate (RGR), net assimilation rate (NAR), height, basal diameter, root length, leaf number, root mass/total plant mass (RMR) and root-to-shoot dry biomass (R/S ratio), and decreased in leaf area ratio (LAR), specific leaf area (SLA), leaf size, stem mass-to-total plant mass ratio (SMR) and leaf mass-to-total plant mass ratio (LMR) with increasing light availability. Under the three light environments, the two types of species differed significantly in total biomass, total leaf area, RGR, NAR, LAR, SLA and leaf number, and not in leaf area. Only light-demanding species differed significantly in height, root length, basal diameter, RMR, SMR, LMR and R/S ratio. The mean plasticity index of growth and biomass allocation were relatively higher than the morphological variables, with significant differences between the two groups. Our results showed that liana species respond differently to changing light environments and that light-demanding species exhibit higher plasticity. Such differences may affect the relative success of liana species in forest dynamics.     

荆条叶性状对野外不同光环境的表型可塑性

光照是影响植物生长和分布的重要环境因子。对生长在野外5种不同光环境下(林外、阔叶林林缘、阔叶林林下、针叶林林窗和针叶林林下)的荆条的叶片进行取样研究,通过对光合作用光响应曲线、叶绿素荧光、叶绿素含量、叶片氮磷含量以及叶片形态的测量,来反映荆条对不同光环境的表型可塑性。研究结果表明,荆条叶片对于野外不同的光环境具有很好的适应机制,叶片功能性状受到结构性状的调节。低光下通过高的比叶面积(SLA)、单位质量叶绿素含量、光系统II最大量子产量,低的暗呼吸速率、光饱和点、光补偿点、叶绿素a,b的比值来提高对光能的利用效率,维持生长;高光下则通过与SLA有关的叶片结构的变化对光合作用进行调节。大多数的叶性状只受到日光照总量的影响,SLA的大小与日最高光强有关,可以对不同日变化模式的光照做出迅速的响应,是适应不同光照的敏感指标。尽管光照是不同光环境下影响荆条叶性状的主要环境因子,土壤养分含量同样会对叶性状产生影响,高土壤养分下的高叶长与叶柄长的比值体现了植物对资源获取和支撑结构之间分配的权衡。     

Interactive effects of clipping and nutrient availability on the compensatory growth of a grass species

Resource availability is an important factor affecting the capacity of compensatory growth after grazing. We performed a greenhouse experiment with Poa bulbosa, a small perennial grass of the Mediterranean and Central Asian grasslands, to test the importance of nutrient availability for compensatory growth after clipping. We also compared the results with predictions of the limited resource model (LRM). Plants were grown at low and high fertilization levels and subjected to a clipping treatment. Contrary to the LMR, we found that in Poa plants compensatory growth occurred under the high fertilization level, while it did not occur under the low level. The LMR predicts a higher tolerance for grazing in the stressful environment. Our plants showed a significant decrease in their relative growth rates (RGR) after clipping. Although the plants allocated a 32–188% greater fraction of the mass to lamina growth after clipping, this greater allocation to the leaves did not fully compensate for the initial reduction in leaf area ratio (LAR). A sensitivity analysis showed for the clipped plants under the high fertilization treatment, that changes in leaf allocation (f _lam) enabled the plants to compensate for a part of the potential loss caused by defoliation. Probably, the increased biomass allocation comes largely from the bulbs. We conclude that the inconsistency of the LRM with our results originates in the lack of compensatory mechanisms in the model. To better understand how environmental conditions affect tolerance to herbivory, the effects of compensatory growth should be taken into account.     

Growth and leaf production in the tropical palm Euterpe edulis: Light conditions versus developmental constraints

We studied developmental and environmental constraints on leaf dynamics, morphology and physiology in the monopodial tropical palm of the Atlantic Forest biome, Euterpe edulis. Plastic responses to light environments in terms of photosynthesis, leaf size, leaf life span, patterns of biomass allocation and growth were analysed. Plants were grown during 14 months in a shade house under four different growth irradiances. Plants of Euterpe edulis were able to adjust leaf demography and biomass allocation in the different light treatments. Leaf life span increased by 100 days with decreasing light levels while the rate of leaf production decreased, consistent with lower electron transport rates. At low light levels, adjustments in biomass allocation to leaf components allowed E. edulis to reduce self-shading and increase light interception. At high light plants allocated more biomass to roots, and the plants exhibited small leaf sizes when leaves were compared using an explicit ontogenetic analysis. Ontogeny constrained the maximum size that each consecutive leaf could achieve, while growth irradiance determined the rate of leaf production and other leaf traits. Consequently, there were both, developmental constraints and environmental determinants influencing leaf demography and morphology in E. edulis. The findings of this ecophysiological and demographic study are relevant to palms growing under natural conditions and help to explain the success of E. edulis in the forest understory and its absence from large gap openings. Our results not only confirm that E. edulis is a shade tolerant species, but also show that palms are able to acclimate to different growing condition as well as trees.     

Combined effects of light and water availability on photosynthesis and growth of Arisaema heterophyllum in the forest understory and an open site

Photosynthetic characteristics, leaf longevity and biomass accumulation of a threatened herb species, Arisaema heterophyllum, were studied in the understory of a riparian forest and at a neighboring deforested open site for 3 years in order to understand the combined effects of light and water availability. Light availability was 2- to 4-fold higher at the deforested than at the forest site during the growing season of the species, and precipitation varied considerably over the 3 years. Despite the difference in water availability among the years (dry in 1994 and 1996, and wet in 1995), the species showed a strong acclimation to the different light environments. Light-saturated assimilation rate on a leaf area basis, leaf mass area ratio (LMA), and relative growth rate (RGR) were all higher at deforested site. While a positive correlation between individual RGR and microsite light availability was found in the wet year, no correlation was found in the dry years, and mean RGR was significantly lower in the dry year at both sites. Leaf longevity, photosynthetic capacity on a leaf mass basis, dark respiration rate, and leaf conductance, varied considerably from year to year, especially in the plants at the open site, probably depending on water availability. In the dry years plants at the deforested sites showed a lower photosynthetic rate and leaf conductance under unwatered than under watered conditions. These results suggest that the water availability in a given year may strongly affect light acclimation and annual RGR of the herb species in natural habitats, even under mesic climate conditions. Received: 15 February 1997 / Accepted: 20 May 1997     

Herbivory on Temperate Rainforest Seedlings in Sun and Shade: Resistance,Tolerance and Habitat Distribution

Differential herbivory and/or differential plant resistance or tolerance in sun and shade environments may influence plant distribution along the light gradient. Embothrium coccineum is one of the few light-demanding tree species in the temperate rainforest of southern South America, and seedlings are frequently attacked by insects and snails. Herbivory may contribute to the exclusion of E. coccineum from the shade if 1) herbivory pressure is greater in the shade, which in turn can result from shade plants being less resistant or from habitat preferences of herbivores, and/or 2) consequences of damage are more detrimental in the shade, i.e., shade plants are less tolerant. We tested this in a field study with naturally established seedlings in treefall gaps (sun) and forest understory (shade) in a temperate rainforest of southern Chile. Seedlings growing in the sun sustained nearly 40% more herbivore damage and displayed half of the specific leaf area than those growing in the shade. A palatability test showed that a generalist snail consumed ten times more leaf area when fed on shade leaves compared to sun leaves, i.e., plant resistance was greater in sun-grown seedlings. Herbivore abundance (total biomass) was two-fold greater in treefall gaps compared to the forest understory. Undamaged seedlings survived better and showed a slightly higher growth rate in the sun. Whereas simulated herbivory in the shade decreased seedling survival and growth by 34% and 19%, respectively, damaged and undamaged seedlings showed similar survival and growth in the sun. Leaf tissue lost to herbivores in the shade appears to be too expensive to replace under the limiting light conditions of forest understory. Following evaluations of herbivore abundance and plant resistance and tolerance in contrasting light environments, we have shown how herbivory on a light-demanding tree species may contribute to its exclusion from shade sites. Thus, in the shaded forest understory, where the seedlings of some tree species are close to their physiological tolerance limit, herbivory could play an important role in plant establishment.     

Small Canopy Gaps Influence Plant Distributions in the Rain Forest Understory1

This study tested three hypotheses regarding how plants respond to the spatial heterogeneity in light availability in the rain forest understory: (1) understory plants occur preferentially in the lighter parts of the understory; (2) under–story palms are more shade tolerant than other understory plants; (3) rain forest plants differ in their ontogenetic response to understory light conditions. The study was carried out in old–growth rain forest in the Yasuní National Park, Amazonian Ecuador. The hypotheses were tested by comparing the distributions of 20 plant species (1454 individuals) over microsites with differing degrees of exposure to canopy gaps to the background distribution of these microsites in the forest. The gap exposure of a given microsite was described by an index based on the number and size of gaps in the canopy to which the site was exposed. Two plant height classes were studied: 0.80–2.49 and 2.50–5 m. The first and third hypotheses were accepted, while the second hypothesis was rejected. The results for the individual species corresponded well with what is known from earlier studies about the ecology of these species or close relatives, suggesting that the patterns observed can be generalized for Neotropical rain forests. Notably, the most abundant species in the study represent several different life history strategies. Thus, abundance in the rain forest understory can be achieved by several different strategies. This suggests that niche differentiation in terms of response to small changes in understory light conditions may be an important factor in the maintenance of the high local plant species richness of tropical rain forests.     

Root/Shoot Allocation and Root Architecture in Seedlings: Variation among Forest Sites, Microhabitats, and Ecological Groups1

I analyzed patterns of variation in root mass allocation and root morphology among seedlings of woody species in relation to environmental factors in four Neotropical forests. Among forests, I explored the response of root traits to sites varying in water or nutrient availability. Within each forest, I explored the plastic response of species to different microhabitats: gaps and understory. Additionally, I explored evidence for life history correlation of root and shoot traits by comparing species differing in their successional group (light‐demanding [22 spp.] or shade tolerant [27 spp.]) and germination type (species with photosynthetic cotyledons or species with reserve cotyledons). At each forest site, young seedlings from 10 to 20 species were excavated. A total of 55 species was collected in understory conditions and 31 of them were also collected in gaps. From each seedling, six morphological ratios were determined. Allocation to roots was higher in forest sites with the lowest soil resources. Roots were finer and longer in the most infertile site, while roots were deeper in the site with the longest dry season. Seedling traits did not differ between germination types. Shade tolerant species allocated more to roots and developed thicker roots than light‐demanding species. Light‐demanding species showed stronger plastic responses to habitat than shade tolerant species, and species with photo‐synthetic cotyledons showed lower plasticity than species with reserve cotyledons. Overall, these results suggest that among Neotropical species, root allocation and root morphology of seedlings reflect plant adjustments to water or nutrient availability at geographic and microhabitat scales. In addition, life history specialization to light environments is suggested by differences among groups of species in their allocation to roots and in their root morphology.     

SEASONAL NUTRIENT DYNAMICS,NUTRIENT RESORPTION,AND MYCORRHIZAL INFECTION INTENSITY OF TWO PERENNIAL FOREST HERBS

Patterns of mycorrhizal infection, seasonal foliar nutrient concentrations, nutrient allocation to shoots and rhizomes, and nutrient resorption were measured in relation to soil nutrient availability in two species of perennial forest herbs, Geranium maculatum L. and Polygonatum pubescens Pursh., in four forest stands in southern and central Ohio. The percentage of plants with V-A mycorrhizae and the proportion of root length colonized by VAM structures increased with decreasing nutrient availability in both species. Foliar N and P concentrations in plants from lower fertility sites were as high, or higher, than those in plants from higher fertility sites; as a result, tissue nutrient enrichment ratios (foliar concentration/soil available concentration) increased with decreasing fertility. Proportional resorption of N and P generally decreased with decreasing nutrient availability, a pattern inconsistent with those exhibited by woody plants in these forest stands. We hypothesize that the inverse relationship between nutrient uptake efficiency (via mycorrhizae) and nutrient use efficiency (resorption) exhibited by these forest understory herbs, but not by trees or herbs from high-light environments, may be related to low-light limitation of energy reserves in the forest understory.     

Trait convergence and plasticity among native and invasive species in resource-poor environments

? Premise of study: Functional trait comparisons provide a framework with which to assess invasion and invasion resistance. However, recent studies have found evidence for both trait convergence and divergence among coexisting dominant native and invasive species. Few studies have assessed how multiple stresses constrain trait values and plasticity, and no study has included direct measurements of nutrient conservation traits, which are critical to plants growing in low-resource environments. ? Methods: We evaluated how nutrient and water stresses affect growth and allocation, water potential and gas exchange, and nitrogen (N) allocation and use traits among a suite of six codominant species from the Intermountain West to determine trait values and plasticity. In the greenhouse, we grew our species under a full factorial combination of high and low N and water availability. We measured relative growth rate (RGR) and its components, total biomass, biomass allocation, midday water potential, photosynthetic rate, water-use efficiency (WUE), green leaf N, senesced leaf N, total N pools, N productivity, and photosynthetic N use efficiency. ? Key results: Overall, soil water availability constrained plant responses to N availability and was the major driver of plant trait variation in our analysis. Drought decreased plant biomass and RGR, limited N conservation, and led to increased WUE. For most traits, native and nonnative species were similarly plastic. ? Conclusions: Our data suggest native and invasive biomass dominants may converge on functionally similar traits and demonstrate comparable ability to respond to changes in resource availability.     

Growth responses to arbuscular mycorrhizae by rain forest seedlings vary with light intensity and tree species

Light intensity and root colonization by arbuscular mycorrhizal (AM) fungi are considered important factors affecting the performance of rain forest plants, yet few studies have examined how these two factors interact. Whether AM colonization promoted growth or caused shifts in biomass allocation in seedlings of four species of Australian rain forest tree (Flindersia brayleana, Acmena resa, Cryptocarya mackinnoniana and Cryptocarya angulata), grown in a glasshouse under light conditions that mimicked the shaded understory (3% PAR) and small light gaps (10% PAR), was examined. Seedlings were grown in sterilized field soil and either inoculated with AM fungi or provided sterile inoculum. Four major findings emerged. First, in all species, seedlings grown in small gap light intensities were larger than seedlings grown in understory light intensities. Second, when seedling biomass was included as a covariate, variation in light intensity was associated with significant shifts in biomass allocation. In all species, leaf area ratio was lower at 10% PAR than at 3% PAR, while root-to-shoot ratio showed the opposite pattern in one of the four species (C. mackinonniana). Third, although percentage root length colonized by AM fungi was greater at 10% PAR than 3% PAR in all species, this difference could be accounted for by variation in seedling size in all species except C. angulata. Fourth, growth and biomass allocation responses to AM colonization varied with light intensity and plant species. AM colonization promoted growth in both light regimes only in F. brayleana, while it had no effect on growth in C. mackinnoniana and C. angulata in either light regime and promoted growth only under high light in A. resa. AM colonization had no effect on leaf area ratio or root-to-shoot ratio in any of the species, and significantly altered specific root length in only one of the four species (C. mackinnoniana). These findings suggest that rain forest seedlings are highly variable in their growth responses to AM colonization and that some of this variability is related to the light intensity of the environment. Given that seedlings may spend many years in the shaded understory, these differences among species could have important effects on long-term seedling performance and seedling community dynamics.     

Growth, Allocation and Water Relations of Shade-grownQuercus rubraL. Saplings Exposed to a Late-season Canopy Gap

For understory saplings to exploit canopy gaps successfully,carbon gain must increase in the gap environment. We predictedthat total biomass of shade-grown red oak saplings would increaseafter exposure to a late-season canopy gap, and that increasedwater and nutrient demand within the canopy gap would drivechanges in the allocation of this carbon. Shade-grown red oaksaplings acclimated to gaps by increasing biomass during theseason of gap formation and increasing the potential for carbongain in the following summer. Within-season carbon gain didnot result from greater production of leaf area, so it mostlikely arose from higher photosynthetic rates of existing shade-developedfoliage, which may be linked to accumulation of leaf nitrogen.During the season of gap formation, shade-gap plants increasedallocation to storage of total non-structural carbohydrates(TNC), and to root growth. The increase in TNC storage suggeststhat shade-developed saplings exposed to gaps were also primedfor fast growth and carbon gain in the following summer. Theincrease in root growth suggests that higher nutrient and waterdemand drove allocation shifts to enhance the capacity for nutrientand water uptake in the gap. Plant hydraulic conductivity (K_a)of shade-grown plants was limited upon exposure to the gap,possibly because of embolism formation resulting from the abruptincrease in water demand. Greater water potential gradientscompensated for limitations to K_a, allowing saplings to maintainhigh transpiration rates, suggesting that actual water uptakeof shade-gap plants was unaffected by gap exposure. Acclimation; canopy gaps; carbon allocation; hydraulic conductivity; nitrogen allocation; non-structural carbohydrates;Quercus rubra L.; red oak; stomatal conductance; transpiration; water-relations     

Survival and growth of Virola surinamensis yearlings: Water augmentation in gap and understory

Summary Factors affecting seedling Virola surinamensis (Myristicaceae) survival and growth were investigated on Barro Colorado Island, Panama. Seedlings planted 3 months after germination were monitored in treefall gaps and understory using 2.25 ha irrigated and control plots through the first dry season. During the dry season, irrigated plants in gaps increased total leaf area significantly more than did irrigated plants in the shaded understory. Over the same dry season, control plants in gaps and in the shaded understory lost similar amounts of leaf area. Seedlings in understory were suppressed in stem height and biomass in both irrigated and control plots; these measures were greater in gaps and greatest in irrigated gaps (height). Roots were similar in length in all treatments, but greater in biomass in gaps than understory due to greater proliferation of secondary roots in control and irrigated gaps than in control and irrigated understory. This experiment demonstrates both water and light limitation during the first dry season after germination. V. surinamensis seedlings are capable of survival and modest growth of leaf area in the deep shade of the understory in moist locations; they are severely disadvantaged in shaded understory subject to drought, where most seeds fall and most seedlings establish. The broken canopy of a gap allows shoot and consequently root growth that permits seedlings to survive seasonal drought.     

不同生境朝鲜淫羊藿生长与光合特征

选择朝鲜淫羊藿野外生存的林缘、林窗和林下3种生境,研究朝鲜淫羊藿形态特征、光合特征、叶绿素含量、生物量积累及分配对不同光环境的响应。结果表明,朝鲜淫羊藿株高、茎径、总叶面积等形态指标随光照强度减弱明显减小,各生境间差异显著。随光照强度降低,朝鲜淫羊藿的净光合速率、水分利用效率、气孔限制值、叶绿素a/b降低,而胞间CO2浓度、光能利用效率、叶绿素b和总叶绿素含量增加。各器官生物量积累随光照强度减弱而降低,各生境间差异极显著,其中,林缘地上生物量是林窗的1.6倍,林下的3.1倍。生物量分配方面,叶生物量比、叶面积比率、比叶面积随光照强度减弱而增加,根状茎生物量比和根冠比降低,林缘、林窗与林下差异极显著;林下、林缘、林窗朝鲜淫羊藿地上部分生物量比分别为0.733、0.659、0.664,林下朝鲜淫羊藿与林缘、林窗存在不同的光利用策略和生存策略。朝鲜淫羊藿作为极易遭受利用和生境破坏威胁的物种,宜在适度荫闭的林缘和疏林下开展野生抚育或仿生态栽培。