Responses of temperate woody seedlings to shade and drought: do trade-offs limit potential niche differentiation?

Diversity in seedling responses to combined shade and drought can drive species niche differences, and thus natural forest and scrub establishment dynamics and diversity. However, inherent trade-offs between shade tolerance and drought tolerance, as hypothesized in the literature, would constrain potential niche differentiation. For thirteen species of European trees and shrubs the impacts were determined of moderate and extreme drought on the growth and survival of first-year seedlings in a given soil volume, in irradiances in the range typical for temperate forest and scrub understories (3% daylight) and large gaps (30% daylight). Pots were watered differentially each second day to equalize soil water content across species and irradiances. Comparisons within and across species supported independent tolerances of shade and drought rather than trade-offs. For all species, drought reduced relative growth rate in dry mass by the same proportion in 3% daylight and in 30% daylight. Consequently, drought generally reduced final dry mass significantly more strongly in 30% daylight than in 3% daylight. Extreme drought led to earlier mortality in 30% daylight than in 3% daylight for nine of the eleven species tested, to earlier mortality in 3% daylight for one species, and to equal mortality rate in both irradiances for one species. For each species, growth-based shade tolerance was quantified as the ratio of absolute growth in low irradiance to that in high irradiance, and growth-based drought tolerance as the ratio of absolute growth in low water supply to that in high water supply. Across species, growth-based shade tolerance varied ≈9-fold, and growth-based drought tolerance ≈2-fold; species’ tolerances correlated respectively with indices of field establishment in shade and drought. Growth-based drought tolerance correlated with survival time in extreme drought. Experimentally quantified shade and drought tolerances varied independently for the 13 species tested, indicating the potential for extensive species niche differentiation in combinations of irradiance and water supply.     

The Effects of Drought and Shade on the Performance,Morphology and Physiology of Ghanaian Tree Species

In tropical forests light and water availability are the most important factors for seedling growth and survival but an increasing frequency of drought may affect tree regeneration. One central question is whether drought and shade have interactive effects on seedling growth and survival. Here, we present results of a greenhouse experiment, in which seedlings of 10 Ghanaian tree species were exposed to combinations of strong seasonal drought (continuous watering versus withholding water for nine weeks) and shade (5% irradiance versus 20% irradiance). We evaluated the effects of drought and shade on seedling survival and growth and plasticity of 11 underlying traits related to biomass allocation, morphology and physiology. Seedling survival under dry conditions was higher in shade than in high light, thus providing support for the “facilitation hypothesis” that shade enhances plant performance through improved microclimatic conditions, and rejecting the trade-off hypothesis that drought should have stronger impact in shade because of reduced root investment. Shaded plants had low biomass fraction in roots, in line with the trade-off hypothesis, but they compensated for this with a higher specific root length (i.e., root length per unit root mass), resulting in a similar root length per plant mass and, hence, similar water uptake capacity as high-light plants. The majority (60%) of traits studied responded independently to drought and shade, indicating that within species shade- and drought tolerances are not in trade-off, but largely uncoupled. When individual species responses were analysed, then for most of the traits only one to three species showed significant interactive effects between drought and shade. The uncoupled response of most species to drought and shade should provide ample opportunity for niche differentiation and species coexistence under a range of water and light conditions. Overall our greenhouse results suggest that, in the absence of root competition shaded tropical forest tree seedlings may be able to survive prolonged drought.     

Photoinhibition and drought in Mediterranean woody saplings: scaling effects and interactions in sun and shade phenotypes

Interacting effects of high light and drought on the performance of sun and shade phenotypes were experimentally undertaken following survival, chlorophyll fluorescence and gas exchange in 2-year-old saplings of four Mediterranean trees (Quercus ilex and Q. coccifera as water-saving species, and Pistacia lentiscus and P. terebinthus as water-spending species). Half of the saplings were grown in full sunlight and the other half in the shade (6% sunlight). Half of each combination of species-phenotype was exposed to high light during a simulated late-summer drought. Light absorptance and gas exchange were scaled up to the whole plant with the 3-D geometrical model, Y-Plant. Quercus species were more plastic and tolerated high light and water stress better than Pistacia species, surviving longer and in drier soils, and exhibiting a less pronounced photoinhibition. There was no evidence of disadvantage for shade phenotypes under high light with increasing drought. By contrast, shade phenotypes survived longer despite larger initial decreases in photochemical efficiency and higher sensitivity to drought than sun phenotypes. The enhanced control of transpiration during drought in water-saving versus water-spending species (and also in shade versus sun phenotypes in three out of the four species) allowed extended survival. Photoinhibition reduced whole crown carbon gain in high light by c. 3% and affected significantly more the shaded leaves of a given plant (reducing their carbon gain by up to 7%) than those exposed to direct sunlight. Despite this apparently minor impact, whole plant carbon gain reduction by photoinhibition negatively correlated with survival and drought tolerance. The implications for succession and forest regeneration in arid environments, particularly under a global change scenario, are discussed.     

Non-linear effects of drought under shade: reconciling physiological and ecological models in plant communities

The combined effects of shade and drought on plant performance and the implications for species interactions are highly debated in plant ecology. Empirical evidence for positive and negative effects of shade on the performance of plants under dry conditions supports two contrasting theoretical models about the role of shade under dry conditions: the trade-off and the facilitation hypotheses. We performed a meta-analysis of field and greenhouse studies evaluating the effects of drought at two or more irradiance levels on nine response variables describing plant physiological condition, growth, and survival. We explored differences in plant response across plant functional types, ecosystem types and methodological approaches. The data were best fit using quadratic models indicating a humped-back shape response to drought along an irradiance gradient for survival, whole plant biomass, maximum photosynthetic capacity, stomatal conductance and maximal photochemical efficiency. Drought effects were ameliorated at intermediate irradiance, becoming more severe at higher or lower light levels. This general pattern was maintained when controlling for potential variations in the strength of the drought treatment among light levels. Our quantitative meta-analysis indicates that dense shade ameliorates drought especially among drought-intolerant and shade-tolerant species. Wet tropical species showed larger negative effects of drought with increasing irradiance than semiarid and cold temperate species. Non-linear responses to irradiance were stronger under field conditions than under controlled greenhouse conditions. Non-linear responses to drought along the irradiance gradient reconciliate opposing views in plant ecology, indicating that facilitation is more likely within certain range of environmental conditions, fading under deep shade, especially for drought-tolerant species.     

Relative importance of photosynthetic traits and allocation patterns as correlates of seedling shade tolerance of 13 tropical trees

Among 13 tropical tree species on Barro Colorado Island, species with high seedling mortality rates during the first year in shade had higher reltive growth rates (RGR) from germination to 2 months in both sun (23% full sun) and shade [2%, with and without lowered red: far red (R:FR) ratio] than shade tolerant species. Species with higher RGR in sun also had higher RGR in shade. These interspecific trends could be explained by differences in morphological traits and allocation paterns among species. Within each light regime, seedlings of shade-intolerant species had lower root: shoot ratios, higher leaf mass per unit area, and higher leaf area ratios (LAR) than shade tolerant species. In contrast, leaf gas exchange characteristics, or acclimation potential in these traits, had no relationship with seedling mortality rates in shade. In both shade tolerant and intolerant species, light saturated photosynthesis rates, dark respiration, and light compensation points were higher for sungrown seedlings than for shade-grown seedlings. Differences in R:FR ratio in shade did not affect gas exchange, allocation patterns, or growth rates of any species. Survival of young tree seedlings in shade did not depend on higher net photosynthesis or biomass accumulation rates in shade. Rather, species with higher RGR died faster in shade than species with lower RGR. This trend could be explained if survival depends on morphological characteristics likely to enhance defense against herbivores and pathogens, such as dense and tough leaves, a well-established root system, and high wood density. High construction costs for these traits, and low LAR as a consequence of these traits, should result in lower rates of whole-plant carbon gain and RGR for shade tolerant species than shade-intolerant species in shade as well as in sun.     

Light-dependent changes in the relationship between seed mass and seedling traits: a meta-analysis for rain forest tree species

Seed mass is considered to be an important attribute for the establishment success of plant species being linked with their seed production, establishment, and survival. This meta-analysis shows that seed mass is also closely correlated to growth-related species attributes of the established phase of rain forest tree species, and that the strength of this relationship varies with light conditions. Seed mass is an especially good predictor of species traits under high-light conditions, when the species attain their full growth potential. At high irradiance (>20% of full light) seed mass is negatively correlated with RGR, NAR, LAR, SLA and LMF. At low irradiance (<5% of full light), seed mass is only negatively correlated with LAR and SLA. Correlations between seed mass and morphological traits are therefore strongest at low irradiance where light interception is important. Conversely, correlations between seed mass and a physiological trait are strongest at high irradiance, where maximisation of photosynthetic rates is important. The strength of the correlation between growth parameters and seed mass declines over time, and disappears after 1–4 years. Seed mass appears to be a good proxy for the shade tolerance of tropical tree species, especially at the younger stages of the life cycle.     

光照对鄂东南2种落叶阔叶树种幼苗生长、光合特性和生物量分配的影响

通过设置4个光照梯度(25%、12%、6%和3%自然光)模拟鄂东南低山丘陵地区落叶阔叶林林下的光环境,研究了2种耐荫性不同的树种幼苗--麻栎(Quercus acutissima)和化香(Platycarya strobilacea)不同光强下的存活率、光合特性、生长和生物量分配,探讨了低光环境中耐荫性不同的树种幼苗维持自身碳平衡的机制和权衡"存活-生长"选择的生活史策略。结果表明:(1)低光下的2个树种幼苗的生长、光合特性和生物量分配具有显著性差异。(2)各个光照梯度下麻栎幼苗都生长良好,存活率保持在35%以上,而化香幼苗遭遇高的死亡率,80d后3%和6%自然光下的幼苗全部死亡;低光环境中麻栎幼苗比化香幼苗具有更大的表观光量子(AQY)和最大净光合效率(Pmax),更低的光补偿点(LCP)和暗呼吸效率(Rd),即耐荫性较强的麻栎幼苗比耐荫性较弱的化香幼苗具有更高的低光碳同化率和碳捕获能力。(3)2个树种幼苗的成活率与RGR呈负相关关系,各个光照梯度下耐荫性较弱的化香幼苗的相对生长率(RGR)显著高于耐荫性较强的麻栎幼苗,而两个树种幼苗的净同化率(NAR)无明显差异。相对于麻栎幼苗较高的根生物量比(RMR),化香幼苗将更多的生物量分配给叶部,因而具有较高的叶生物量比(LMR)、叶面积比(LAR)和比叶面积(SLA)。不同耐荫性的幼苗生长及生物量分配方式的差异是植物"存活-生长"权衡后的结果,耐荫性弱的化香幼苗具有较高的生长潜力和较弱的自我保护能力,而耐荫性强的麻栎幼苗具有更高的低光碳储量,能够维持更好的低光碳平衡,具有竞争优势。     

Is hemiepiphytism an adaptation to high irradiance? Testing seedling responses to light levels and drought in hemiepiphytic and non‐hemiepiphytic Ficus

The epiphytic growth habit in many Ficus species during their juvenile stages has commonly been hypothesized to be an adaptation for avoiding deep shade in the forest understory, but this has never been tested experimentally. We examined growth and ecophysiology in seedlings of three hemiepiphytic (Hs) and three non‐hemiepiphytic (NHs) Ficus species grown under different irradiance levels. Both Hs and NHs exhibited characteristics of high light requiring species, such as high plasticity to growth irradiance and relatively high maximum photosynthetic assimilation rates. Diurnal measurements of leaf gas exchange showed that Hs have much shorter active photosynthetic periods than NHs; moreover, leaves of Hs have lower xylem hydraulic conductivity but stronger drought tolerance as indicated by much lower rates of leaf diebacks during the drought treatment. Seedlings of NHs had 3.3‐ and 13.3‐fold greater height and biomass than those of Hs species after growing in the nursery for 5 months, indicating a trade‐off between growth and drought tolerance due to the conflicting requirements for xylem conductivity and cavitation resistance. This study does not support the shade‐avoidance hypothesis; rather, it suggests that the canopy regeneration in Hs is an adaptation to avoid alternative terrestrial growth‐related risks imposed to tiny Ficus seedlings. The NHs with terrestrial regeneration reduce these risks by having an initial burst of growth to rapidly gain relatively large seedling sizes, while in Hs seedlings more conservative water use and greater drought tolerance for surviving the canopy environment are intrinsically associated with slow growth.     

Tertiary relict trees in a Mediterranean climate: abiotic constraints on the persistence of Prunus lusitanica at the eroding edge of its range

Aim To investigate the ecophysiological traits allowing persistence of a subtropical relict tree (Prunus lusitanica L.) under a dry Mediterranean climate at the eroding edge of its range. Location A glasshouse for the study under controlled conditions and two marginal populations located in riparian forests of central Spain and exposed to summer drought, in contrast to subtropical populations that grow in mountain cloud forests. Methods Two experiments were conducted to assess tolerance to the abiotic conditions found in riparian habitats. In a glasshouse experiment, gas‐exchange and light‐use parameters were measured to evaluate seedling responses to a factorial combination of irradiance (60%, 10%, 2% and 0.5% full sun) and moisture (well watered control and drought stress). In a parallel field experiment, irradiance and soil moisture were measured as predictors of seedling survival at two sites in which half the seedlings were subjected to a threefold increase in summer precipitation by adding water every 10 days. Results  Soil moisture strongly determined survival both in the glasshouse and in the two field sites. In the field, periodic addition of water failed to increase survival. Water‐use efficiency (WUE) increased with drought. Seedlings did not tolerate deep shade (2–0.5%) and their performance and survival were better when exposed to moderate (10%) or high (60%) irradiance. The effect of water stress on seedling performance was stronger at moderate irradiance, moderate at high irradiance and negligible at very low light. Seedling size (height and number of leaves) attained after 1 month of experimental treatments had a positive effect on survival at the end of the summer, hence greater survival was not achieved at the expense of vertical growth. Main conclusions While studies in Macaronesia have shown that P. lusitanica occupies a wide range of moisture conditions along mountain slopes, it behaves as an obligate riparian species in dry peripheral populations. Intolerance to deep shade and tolerance to moderate and high irradiance allow the species to grow in small and moist gaps, or in treeless river sections. Despite the species’ low range filling in marginal, drought‐prone regions, long‐term persistence might have been achieved through shifts to riparian habitats serving as local refuges.     

Seed germination and seedling performance of two Mediterranean tree species,holm oak (shape Quercus ilex L.) and Aleppo pine (shape Pinus halepensis Mill.): a multifactor experimental approach

A two-level multifactor experimental approach was used to compare seed germination and seedling performance of two Mediterranean tree species: the early successional Aleppo pine (Pinus halepensis Mill.) and the late successional holm oak (Quercus ilex L.). In a first experiment germination rate was evaluated under the combined effects of shade, nitrogen availability, and pine or holm oak leaf litter. In a second experiment we tested for the effects of shade, nutrient availability, and litter type on seedling survival, growth and biomass allocation. Holm oak showed higher germination rates under shaded than under unshaded conditions, while Aleppo pine showed no differences between shaded and unshaded conditions. Nitrogen availability and litter type had no significant effect on germination of either species. Both species showed increased RGR, but also higher mortality rates, when grown in an enriched nutrient environment. While Aleppo pine showed no differences in RGR and mortality rate under different shading levels, RGR decreased and mortality increased for holm oak in full light. Increased radiation decreased LAR, SLA and height:diameter ratio, and increased RWR and R/S in both species, although Aleppo pine showed more pronounced changes. Unlike Aleppo pine, holm oak responded to increased nutrient availability by decreasing R/S and increasing LAR. From these results, no seed-seedling conflicts were found in either species, but a trade-off does seem to exist for holm oak between biomass allocation traits deployed in response to increased nutrient availability and radiation. Aleppo pine outperformed holm oak under most environmental conditions tested and showed a wider regeneration niche.     

Shade tolerance and herbivory are associated with RGR of tree species via different functional traits

• Relative growth rate (RGR) plays an important role in plant adaptation to the light environment through the growth potential/survival trade‐off. RGR is a complex trait with physiological and biomass allocation components. It has been argued that herbivory may influence the evolution of plant strategies to cope with the light environment, but little is known about the relation between susceptibility to herbivores and growth‐related functional traits.
• Here, we examined in 11 evergreen tree species from a temperate rainforest the association between growth‐related functional traits and (i) species’ shade‐tolerance, and (ii) herbivory rate in the field. We aimed at elucidating the differential linkage of shade and herbivory with RGR via growth‐related functional traits.
• We found that RGR was associated negatively with shade‐tolerance and positively with herbivory rate. However, herbivory rate and shade‐tolerance were not significantly related. RGR was determined mainly by photosynthetic rate (A_max) and specific leaf area (SLA). Results suggest that shade tolerance and herbivore resistance do not covary with the same functional traits. Whereas shade‐tolerance was strongly related to A_max and to a lesser extent to leaf mass ratio (LMR) and dark respiration (R_d), herbivory rate was closely related to allocation traits (SLA and LMR) and slightly associated with protein content.
• The effects of low light on RGR would be mediated by A_max, while the effects of herbivory on RGR would be mediated by SLA. Our findings suggest that shade and herbivores may differentially contribute to shape RGR of tree species through their effects on different resource‐uptake functional traits.

     

Interactions of drought and shade effects on seedlings of four Quercus species: physiological and structural leaf responses

Here, we investigated the physiological and structural leaf responses of seedlings of two evergreen and two deciduous Quercus species, grown in a glasshouse and subjected to contrasted conditions of light (low, medium and high irradiance) and water (continuous watering vs 2-months drought). The impact of drought on photosynthetic rate was strongest in high irradiance, while the impact of shade on photosynthetic rate was strongest with high water supply, contradicting the hypothesis of allocation trade-off. Multivariate causal models were evaluated using d-sep method. The model that best fitted the dataset proposed that the variation in specific leaf area affects photosynthetic rate and leaf nitrogen concentration, and this trait determines stomatal conductance, which also affects photosynthetic rate. Shade conditions seemed to ameliorate, or at least not aggravate, the drought impact on oak seedlings, therefore, the drought response on leaf performance depended on the light environment.     

Functional analysis of the relative growth rate, chemical composition, construction and maintenance costs, and the payback time of Coffea arabica L. leaves in response to light and water availability

In this study, the combined effects of light and water availability on the functional relationships of the relative growth rate (RGR), leaf chemical composition, construction and maintenance costs, and benefits in terms of payback time for Coffea arabica are presented. Coffee plants were grown for 8 months in 100% or 15% full sunlight and then a four-month water shortage was implemented. Plants grown under full sunlight were also transferred to shade and vice versa. Overall, most of the traits assessed were much more responsive to the availability of light than to the water supply. Larger construction costs (12%), primarily associated with elevated phenol and alkaloid pools, were found under full sunlight. There was a positive correlation between these compounds and the RGR, the mass-based net carbon assimilation rate and the carbon isotope composition ratio, which, in turn, correlated negatively with the specific leaf area. The payback time was remarkably lower in the sun than in shade leaves and increased greatly in water-deprived plants. The differences in maintenance costs among the treatments were narrow, with no significant impact on the RGR, and there was no apparent trade-off in resource allocation between growth and defence. The current irradiance during leaf bud formation affected both the specific leaf area and leaf physiology upon transferring the plants from low to high light and vice versa. In summary, sun-grown plants fixed more carbon for growth and secondary metabolism, with the net effect of an increased RGR.     

Trade-offs between seedling growth and survival in deciduous broadleaved trees in a temperate forest

BACKGROUND AND AIMS: In spatially heterogeneous environments, a trade-off between seedling survival and relative growth rate may promote the coexistence of plant species. In temperate forests, however, little support for this hypothesis has been found under field conditions, as compared with shade-house experiments. Performance trade-offs were examined over a large resource gradient in a temperate hardwood forest. METHODS: The relationship between seedling survival and seedling relative growth rate in mass (RGR(M)) or height (RGR(H)) was examined at three levels of canopy cover (forest understorey, FU; small gap, SG; and large gap, LG) and at two microsites within each level of canopy cover (presence or absence of leaf litter) for five deciduous broad-leaved tree species with different seed sizes. KEY RESULTS: Within each species, both RGR(M) and RGR(H) usually increased with increasing light levels (in the order FU < SG < LG), whereas little difference was observed based on the presence or absence of litter. Seedling survival in FU was negatively correlated with both RGR(M) and RGR(H) in both LG and SG. The trade-off between high-light growth and low-light survival was more evident in the relationship with LG as compared with SG. An intraspecific trade-off between survival and RGR was observed along environmental gradients in Acer mono, whereas seedlings of Betula platyphylla var. japonica survived and grew better in LG. CONCLUSIONS: The results presented here strongly support the idea of light gradient partitioning (i.e. species coexistence) in spatially heterogeneous light environments in temperate forests, and that further species diversity would be promoted by increased spatial heterogeneity. The intraspecific trade-off between survival and RGR in Acer suggests that it has broad habitat requirements, whereas Betula has narrow habitat requirements and specializes in high-light environments.     

Patterns of dynamic irradiance affect the photosynthetic capacity and growth of dipterocarp tree seedlings

In the deeply shaded understorey of S.E. Asian rain forests the growth and survival of dipterocarp seedlings is limited by their ability to maintain a positive carbon balance. Photosynthesis during sunflecks is an important component of carbon gain in understorey plants. To test the sensitivity of photosynthesis and growth to variation in the pattern of dynamic irradiance, dipterocarp tree seedlings (Shorea leprosula and Hopea nervosa) were grown for 370 days under shaded forest light treatments of equal total daily photosynthetic photon flux density (approximately 3.3 mol m(-2) day(-1)), but characterised by either long flecks (LF) or short flecks (SF). Seedling growth was more than 4-fold greater under LF, compared with SF, in both species. Variation in the relative growth rates (RGR) and light saturated rates of photosynthesis (A(max)) were strongly positively correlated with the mean duration of sunflecks. Variation in RGR was strongly correlated with greater unit leaf rate growth, indicating that photosynthetic carbon gain per unit leaf area was greater under LF. The accumulation of starch in leaves over the diurnal period was 117% greater in both species under LF, compared with SF. Greater carbon gain in seedlings under LF is likely to have resulted from the combination of (1) greater A(max) (S. leprosula 35%, H. nervosa 40%), (2) more efficient dynamic photosynthesis, and (3) greater incident photosynthetic quantum yield, compared with seedlings receiving the SF irradiance treatment. The pattern of dynamic irradiance received by seedlings may significantly impact their growth and survival to a previously unrecognised extent, with important consequences for regeneration processes and hence forest structure and composition.     

Irradiance and temperature affect the competitive interference of blackberry on the physiology of European beech seedlings

The potential negative influence of competition from early successional species like blackberry (Rubus fruticosus) may be decisive for the natural regeneration success of drought-sensitive beech (Fagus sylvatica), especially in the light of climate change. With a split plot glasshouse experiment, we investigated the influence of two air temperature and irradiance levels on the competitive interference of blackberry on the water, nitrogen (N) and carbon (C) balance of beech seedlings under moderate drought. When increased temperature was accompanied by low irradiance the biomass, root-to-shoot ratio, N uptake and assimilation rates of blackberry were lower compared with beech, either grown alone or with blackberry. By contrast, when elevated temperature and high irradiance were combined, the root-to-shoot ratio and specific N uptake rate of blackberry were substantially increased, while the N acquisition of beech was impaired. Under lower temperature, with either full light or shade, the presence of blackberry had no significant effects on beech, for almost all tested parameters. Under elevated air temperature beech was impaired by the presence of blackberry at high irradiance. These findings emphasize the interacting effects between environmental factors and competition on the establishment of beech regeneration, which should be considered for future forest management in the frame of climate change.     

Trade‐offs in seedling growth and survival within and across tropical forest microhabitats

For niche differences to maintain coexistence of sympatric species, each species must grow and/or survive better than each of the others in at least one set of conditions (i.e., performance trade‐offs). However, the extent of niche differentiation in tropical forests remains highly debated. We present the first test of performance trade‐offs for wild seedlings in a tropical forest. We measured seedling relative growth rate (RGR) and survival of four common native woody species across 18 light, substrate, and topography microhabitats over 2.5 years within Hawaiian montane wet forest, an ideal location due to its low species diversity and strong species habitat associations. All six species pairs exhibited significant performance trade‐offs across microhabitats and for RGR versus survival within microhabitats. We also found some evidence of performance equivalence, with species pairs having similar performance in 26% of comparisons across microhabitats. Across species, survival under low light was generally positively associated with RGR under high light. When averaged over all species, topography (slope, aspect, and elevation) explained most of the variation in RGR attributable to microhabitat variables (51–53%) followed by substrate type (35–37%) and light (11–12%). However, the relative effects of microhabitat differed among species and RGR metric (i.e., RGR for height, biomass, or leaf area). These findings indicate that performance trade‐offs among species during regeneration are common in low‐diversity tropical forest, although other mechanisms may better explain the coexistence of species with small performance differences.     

Combined effects of shade and drought on tulip poplar seedlings: trade‐off in tolerance or facilitation?

It has been hypothesised that many natural plant distribution patterns can be explained by a trade-off between shade and drought tolerance which would make plants more sensitive to shade under dry conditions. On the other hand, shading by nurse plants is often observed to enhance growth and survival of smaller plants in dry areas. This paper describes an experiment designed to address the interactive effects of drought and shade on growth and physiology of tulip poplar seedlings ( Liriodendron tulipifera ) grown in shade houses under different combinations of irradiance (1%, 5%, 12%, 17%, and 27% of ambient PAR) and soil water content (5–9%, 11–15%, and>20%). The results show no evidence that higher PAR levels compensate for the negative effects of drought on photosynthesis and growth. Rather, rates of estimated daily shoot carbon gain decreased with PAR under dry conditions. Daily xylem sapflow patterns indicated that this was associated with a strong reduction in stomatal conductance in plants growing in dry soil and high PAR conditions. Whole-shoot light compensation points were higher for plants raised under higher PAR conditions, but were not significantly influenced by the water treatments. Shoot dark respiration rates decreased under drier conditions. These results do not support the hypothesis of a trade-off between shade and drought tolerance. Instead, they indicate mechanisms that can lead to positive effects of shading by neighbouring plants under dry conditions. Indeed, such facilitation is thought to be important in many plant communities.     

The influence of competition from herbaceous vegetation and shade on simulated browsing tolerance of coniferous and deciduous saplings

The ability of saplings to tolerate browsing (i.e. the ability to persist with reduced biomass and to compensate for biomass loss) is influenced by the level of stress and their growth strategies. Ultimately, insight into species‐specific responses of saplings to browsing, shade and competition from neighbours will help explain diversity, structure and function of grazed ecosystems such as the endangered wood‐pasture systems. We measured the survival, whole‐sapling biomass and compensatory growth responses of two coniferous (Picea abies and Abies alba) and two deciduous (Acer pseudoplatanus and Fagus sylvatica) tree species to simulated summer browsing (one single clipping event), shade (installation of a shade cloth) and neighbour removal (mowing surrounding vegetation to ground level) treatments and the interactions between them after two‐growing seasons. For all species, there were interacting effects on growth of browsing and environmental condition (shade and neighbours). Simulated browsing resulted in relatively smaller growth losses when plants were growing slowly due to competitive conditions related to herbaceous neighbours. Although none of the clipped saplings could fully compensate for their biomass losses, the saplings were closer to compensation under high competitive conditions than under low competitive conditions. Survival of the clipped saplings remained relatively high and was only significantly reduced for Picea and Acer. Picea was least tolerant of competition and was the only species for which growth was not negatively affected by strong irradiance of a mountain pasture. Surprisingly, the tolerance of saplings to herbivory as browsing tolerance was enhanced under conditions that negatively affected sapling performance (i.e. survival and growth). Apparently, the relative impact of browsing at the early sapling stage is linked to tree life history characteristics such as competition and shade tolerance and will be lower in situations with intense competitive interactions and/or strong irradiance.     

Osservazioni Preliminari Sull'ecologia di Semenzali di Rovere (Quercus Petraea (Matt.) Liebl.)

Abstract

Preliminary observations on the ecology of sessile oak (Quercus petraea (Matt.) Liebl.) seedlings.—The responses of sessile oak seedlings to four different light intensities and to increasing drought (three soil moisture contents) have been investigated in a nursery experiment in order to improve the knowledge of the ecology of this species in the Mediterranean area. Data have been compared with observations on natural regeneration established in different shade and soil moisture conditions in a sessile oak stand. The results of the two experiments showed that low light intensities (transmittance <5%) did not limit the seedlings establishment but induced lowest growth rates, while in full daylight seedlings mortality was very high (61%). In accordance with the observations in natural conditions, seedlings showed the best performances at intermediate light levels (50–80% of full daylight) and with increasing soil moisture: mortality was very low and average height increment, dry weight, relative growth rate (RGR) and net assimilation rate (NAR) showed the highest values. The differences in specific leaf area (SLA), leaf area ratio (LAR), root/stem ratio enabled to discuss the sessile oak seedlings ecology and adaptations to different light intensities and soil moisture content.