Changes in photosynthesis and leaf characteristics with tree height in five dipterocarp species in a tropical rain forest

Variations in leaf photosynthetic, morphological and biochemical properties with increasing plant height from seedlings to emergent trees were investigated in five dipterocarp species in a Malaysian tropical rain forest. Canopy openness increased significantly with tree height. Photosynthetic properties, such as photosynthetic capacity at light saturation, light compensation point, maximum rate of carboxylation and maximum rate of photosynthetic electron transport, all increased significantly with tree height. Leaf morphological and biochemical traits, such as leaf mass per area, palisade layer thickness, nitrogen concentration per unit area, chlorophyll concentration per unit dry mass and chlorophyll to nitrogen ratio, also changed significantly with tree height. Leaf properties had simple and significant relationships with tree height, with few intra- and interspecies differences. Our results therefore suggest that the photosynthetic capacity of dipterocarp trees depends on tree height, and that the trees adapt to the light environment by adjusting their leaf morphological and biochemical properties. These results should aid in developing models that can accurately estimate carbon dioxide flux and biomass production in tropical rain forests.     

Photosynthesis of a tropical canopy tree, Ceiba pentandra, in a lowland forest in Panama

Diel (24 h) courses of CO(2) and water-vapor exchange of Ceiba pentandra (L.) Gaertn. (Bombacaceae) were studied under natural tropical conditions in the semi-evergreen moist forest of Barro Colorado Island, Panama. Measurements were conducted from early February 1991 (dry season), shortly after new leaves emerged, until mid-October 1991 (wet season), when leaves were shed. Rates of net CO(2) uptake were significantly higher in the dry season than in the wet season, and showed a linear decrease with leaf age. Leaf nitrogen concentrations and contents also decreased with age. Our estimate of annual carbon gain (2640 g CO(2) m(-2) year(-1) or 21 g CO(2) g(DW) (-1) year(-1)) is considerably higher than estimates available for temperate forest trees.     

Relationship between tree canopy height and the production of pasture species in a silvopastoral system based on alder trees

Silvopastoral systems in New Zealand that incorporate trees planted to control soil erosion on hills largely rely on the productivity of the pastoral system for financial returns. The effect on pasture productivity of increasing the tree canopy height by pruning Italian gray alder (Alnus cordata) was investigated by measuring the response of light, soil moisture, soil temperature, pasture production of major pasture species, and grazing behaviour of sheep. A split-plot design with four replicates was used. The main plot treatments were three levels of shade (81, 23, and 12% of available photosynthetic photon flux (PPF)), created by pruning 11 year old alder grown at the same density. The sub-plot treatments were four pasture mixes: perennial ryegrass (Lolium perenne), Yorkshire fog (Holcus lanatus), and cocksfoot (Dactylis glomerata), each sown with white clover (Trifolium repens), and cocksfoot sown with lotus (Lotus pedunculatus). Soil temperature was highest under light shade. Total herbage yield at 50 mm stubble height from October to May under heavy and medium shade was 60 and 80%, respectively, of the total herbage harvested under light shade. Cocksfoot had the greatest herbage yield, either with lotus or white clover. The tillering of perennial ryegrass was suppressed by shade more than for the other grass species making ryegrass unsuitable for use in this silvopastoral system. More sheep grazed in the light shade than in the heavy shade, but there was no difference in sheep preference for cocksfoot or Yorkshire fog. Lotus was grazed more frequently than white clover. Pruning of alder to increase canopy height has the potential to improve the productivity of the understorey pasture and its acceptability to sheep.     

Density-dependent selfing and its effects on seed performance in a tropical canopy tree species, Shorea acuminata (Dipterocarpaceae)

In the conservation and management practices of natural forests, sound reproduction and regeneration form the basis of the maintenance and viability of the tree populations. To obtain and serve biological information for sustainable forest management, we investigated reproductive biology and inbreeding depression in seeds of an important dipterocarp tree species, Shorea acuminata (Dipterocarpaceae), by both field and laboratory experiments. Results of parental analysis of immature and mature seeds showed that selfing rates varied greatly, from 7.6 to 88.4% among eight mother trees, and the mean overall selfing rate was 38.3%. Observed outcrossing events within a 40-ha study plot were predominantly (76.5%) short-distance events with a mating distance (md) ≤ 100 m. Since the selfing rate sharply decreased with increase in the number of flowering conspecifics (i.e., individuals of the same species) within a 100-m radius from the mother trees, the local density of flowering conspecifics appears to be the key factor determining the outcrossing rate in S. acuminata. However, the extremely high selfing rate (88.4%) observed for one tree could not be simply explained by the low local density of flowering conspecifics. Instead, differences in its flowering phenology (its flowering peaked ca. a week earlier than most of the other examined individuals) may have severely limited its receipt of pollen from other conspecifics, and thus promoted selfing. Since there were no significant differences in the proportion of selfed progeny between immature and mature seed stages, there was no evidence of selective abortion of selfed seeds during seed development. However, the seed mass of outcrossed progeny was heavier than that of selfed progeny, and heavier seeds showed higher success rates at germination and seedling establishment. These results suggest that inbreeding depression resulted in reductions in seed mass and may reduce the fitness of selfed seeds in S. acuminata. In addition, the outcrossing rate of S. acuminata was more sensitive to low local conspecific flowering-tree densities than that of a sympatric bee-pollinated dipterocarp species with greater pollination distances. These results suggest that the management of local adult-tree densities is important for avoiding selfing and inbreeding depression in future generations, especially in a species like S. acuminata with predominantly short-distance pollination.     

Changes in sapwood permeability and anatomy with tree age and height in the broad-leaved evergreen species Eucalyptus regnans

Increases in plant size and structural complexity with increasing age have important implications for water flow through trees. Water supply to the crown is influenced by both the cross-sectional area and the permeability of sapwood. It has been hypothesized that hydraulic conductivity within sapwood increases with age. We investigated changes in sapwood permeability (k) and anatomy with tree age and height in the broad-leaved evergreen species Eucalyptus regnans F. Muell. Sapwood was sampled at breast height from trees ranging from 8 to 240 years old, and at three height positions on the main stem of 8-year-old trees. Variation in k was not significant among sampling height positions in young trees. However, k at breast height increased with tree age. This was related to increases in both vessel frequency and vessel diameter, resulting in a greater proportion of sapwood being occupied by vessel lumina. Sapwood hydraulic conductivity (the product of k and sapwood area) also increased with increasing tree age. However, at the stand level, there was a decrease in forest sapwood hydraulic conductivity with increasing stand age, because of a decrease in the number of trees per hectare. Across all ages, there were significant relationships between k and anatomy, with individual anatomical characteristics explaining 33-62% of the variation in k. There was also strong agreement between measured k and permeability predicted by the Hagen-Poiseuille equation. The results support the hypothesis of an increase in sapwood permeability at breast height with age. Further measurements are required to confirm this result at other height positions in older trees. The significance of tree-level changes in sapwood permeability for stand-level water relations is discussed.     

Specific leaf area for five tropical tree species growing in different tree species mixtures in Central Panama

New Forests - Leaf traits have been shown to explain a great fraction of differences in growth rates in trees. With this study, we evaluate if differences in growth performance of different species...     

A link between hurricane-induced tree sprouting, high stem density and short canopy in tropical dry forest

The physiognomy of Caribbean dry forest is shorter, denser and contains a greater proportion of multi-stemmed trees than other neotropical dry forests. Our previous research, conducted after Hurricane Georges in 1998, has shown that dry forest trees sprout near the base following hurricane disturbance, even if the trees have not incurred structural damage. However, for these hurricane-induced sprouts to contribute to the physiognomy of the forest, they must grow and survive. We followed sprout dynamics and stem mortality on 1,407 stems from 1998, after Hurricane Georges, until 2005. The number of surviving sprouts and the proportion of sprouting stems decreased during the 7-year period, but the sprouting rate was still 3-fold higher and the proportion of sprouting stems 5-fold higher than before the hurricane. Mortality of non-sprouting stems (15.4%) was about the same as for sprouting stems (13.9%) after 7 years. The mean length of the dominant sprout surpassed 1.6 m by 2005, with over 13% of the dominant sprouts reaching subcanopy height. Sprout growth and survival varied among species. These results demonstrate that, despite some thinning, hurricane-induced sprouts survive and grow and that the unique physiognomic characteristic of Caribbean dry forests is related to hurricane disturbance.     

Optimality and nitrogen allocation in a tree canopy

Physical and functional properties of foliage were measured at a variety of microsites in a broad-leaved Nothofagus fusca (Hook. f.) ?rst. canopy. The light climate of the foliage at these sites was monitored for 39 days in the late spring and early summer with in situ sensors. Foliage nitrogen content (N), mean leaf angle, and gas exchange characteristics were all correlated with the amount of light reaching the microsites during foliage development. Foliage N content on a leaf area basis ranged between ~1 and 2.5 g N m(-2) and was highest at the brightest sites. Light-saturated photosynthetic rates ranged between ~4 and 9 micro mol m(-2) s(-1), increasing from the darkest to brightest sites. A biochemical model of photosynthesis was fitted to foliage characteristics at the different microsites and used to integrate foliage assimilation among the sites over 39 days. The actual arrangement of foliage physiological characteristics in the observed microsites led to higher total canopy rates of net assimilation than > 99% of the combinations of observed foliage characteristics randomly assigned to the observed microsites. Additional simulations first related the maximum rates of electron transport (J(max)), ribulose bisphosphate turnover (V(c,max)), and dark respiration (R(d)) of Nothofagus fusca foliage to nitrogen content and then allowed foliage N (and consequently leaf gas exchange characteristics) to vary across the canopy. The observed N allocation pattern results in greater total canopy assimilation than uniform or > 99% of the simulations with random distributions of N among the microsites (constrained so that the total N allocated was equivalent to that observed in the microsites). However, the observed pattern of N allocation places less N in the brightest microsites and results in substantially less total assimilation than a simulated canopy in which N was allocated in an optimal manner where the N distribution is such that the partial derivative of leaf assimilation (A) with respect to leaf nitrogen content, partial differential A/ partial differential N, is constant among microsites. These results suggest that other factors such as wind or herbivory reduce the integrated assimilation of high-N foliage relatively more than lower-N foliage and that a partial differential A/ partial differential N optimality criteria based only on formulations of leaf gas exchange overestimate canopy assimilation.     

Rank reversals in tree growth along tree size, competition and climatic gradients for four forest canopy dominant species in Central Spain

? Interspecific differences in tree growth patterns with respect to biotic and abiotic factors are key for understanding forest structure and dynamics, and predicting potential changes under climate change.

? Repeated observations from the Spanish Forest Inventory (SFI) were used to parameterize maximum likelihood estimators of tree growth as a function of tree size, competition indices and climate for Pinus pinaster, P. sylvestris, Quercus ilex and Q. pyrenaica.

? Significant responses to both biotic and abiotic factors were found, with interspecific differences in species performance along competition, temperature and precipitation gradients. Q. ilex was the species most tolerant to competition while P. pinaster was the species most sensitive to climatic variation. Species relative positions shifted along gradients of these factors with rank reversals in species performance along size, competition and climatic gradients.

? The results based on average growth matched previous forestry classifications and experimental studies on relative growth rate (RGR).

? When examining growth along studied abiotic and biotic gradients, a mismatch was found between species performance ranks as predicted by our models and information derived from previous knowledge. Those discrepancies highlight the relevance of ontogeny and environmental heterogeneity in defining species performance along competition gradients.

     

Limitation of leaf carbon gain by stomatal and photochemical processes in the top canopy of Macaranga conifera,a tropical pioneer tree

Diurnal changes in gas exchange and chlorophyll fluorescence were measured in the top canopy leaves of the tropical rainforest tree species, Macaranga conifera (Zoll.) Muell. Arg. during a drought year. Maximum values of net photosynthetic rate (P(n), 10 &mgr;mol m(-2) s(-1)) and stomatal conductance (g(s), 0.2 mol m(-2) s(-1)) were found in east-facing leaves in early morning. After 1000 h, both P(n) and g(s) decreased. Minimum daytime values of P(n), g(s), and photosystem II (PSII) quantum yield (DeltaF/F(m)') were found in horizontally fixed leaves. At a given electron transport rate through PSII (ETR), P(n) was higher in early morning than at midday, suggesting a high rate of photorespiration at midday. We tested the hypothesis that the effect of low leaf temperature (T(leaf)) on P(n) is significant in the early morning, whereas the effect of low g(s) on P(n) predominates at midday. In the early morning, when T(leaf) was increased from 32 to 38 degrees C by artificial heating, P(n) at a given ETR decreased 29%, suggesting that the low T(leaf) was associated with a high P(n). When T(leaf) at midday was decreased from 37 to 32 degrees C by artificial cooling, P(n) increased 22%, but P(n) at a given ETR was higher in early morning than at midday, even at the same low T(leaf) (32 degrees C). This suggests that the rate of photorespiration was higher at midday than in early morning because low g(s) at midday caused a reduction in leaf intercellular CO(2) concentration. We conclude that low P(n) at midday was the result of both a reduction in the photochemical process and an increase in stomatal limitation.     

Gender-specific patterns of aboveground allocation, canopy conductance and water use in a dominant riparian tree species: Acer negundo

Acer negundo Sarg. (box elder) is a dioecious tree species that dominates riparian systems at mid elevations throughout the southwest and Intermountain West of the United States. Previous studies have shown that female A. negundo trees occur at higher frequencies along stream margins, whereas males occur at higher frequencies in drier microsites. To better understand the adaptive significance of sex ratio biases and their impact on the ecohydrology of riparian ecosystems, we examined whole-plant water relations and hydraulic properties of mature male and female A. negundo trees occurring within 1 m of a perennial stream channel. We hypothesized that (1) females would have significantly greater canopy water fluxes than males (particularly during periods of seed production: May-June), and (2) xylem in females is more hydraulically efficient but more vulnerable to cavitation than xylem in males. Mean sap flux density (J(s)) during the early growing season (May and June) was 43% higher in female trees than in male trees (n = 6 and 7 trees respectively, P < 0.0001). Mean J(s) in July and August remained 17% higher in females than in males (P = 0.0009). Mean canopy stomatal conductance per unit leaf area (g(s,leaf)) in May and June was on average 140% higher in females than in males (P < 0.0001). Mean g(s,leaf) in July and August remained 69% higher in female trees than in male trees (P < 0.0001). Canopy stomatal conductance scaled to basal area was 90 and 31% higher in females relative to males during May-June and July-August, respectively (P < 0.0001 during both periods). Conversely, there were no apparent differences in either branch hydraulic conductance or branch xylem cavitation vulnerability between genders. These results improve our capacity to describe the adaptive forces that shape the spatial distribution of male and female trees in dioecious species, and their consequences for ecohydrological processes in riparian ecosystems.     

热带雨林濒危树种望天树的致危原因及保护策略

望天树自身的开花结实特点、种子顽拗性和低成苗率、幼苗生长缓慢、过度砍伐和生境破碎化等导致其更新困难,以致陷入濒危的境地。加强自然保护区管理与法制建设、在天然林保护和次生林恢复工作中辅以人工抚育、防治病虫害和人工接种菌根菌以及实施迁地保护等措施可有效地保护望天树种质资源。     

Foliar and shoot allometry of pollarded black locust, Robinia pseudoacacia L.

Browse of multipurpose tree species such as black locust could be used to broaden grazing options, but the temporal distribution of foliage has not been adequately studied. Our objective was to determine effects of harvest date, P fertilization (0 and 600 kg ha⁻¹ yr⁻¹), and pollard height (shoots clipped at 5-, 50-, and 100-cm above ground) on foliar and shoot allometry of black locust. The experiment was conducted on a naturally regenerated 2-yr-old black locust stand (15,000 trees ha⁻¹). Basal shoot diameter and foliar mass were measured monthly in June to October 2002 and 2003. Foliar and shoot dry mass (Y) was estimated from basal shoot diameter (D) by the function Y = aD ^b , with regression explaining ≥95% of variance. Allometry of foliar mass was affected by harvest date, increasing at a greater rate with D in September than in June or July, but not by P fertilization or pollard height. Foliar mass was predicted best by month-specific equations for the June to October growth interval. Allometry of shoot dry mass was unaffected by harvest date, P fertilization, or pollard height. These equations could be used as a first approximation of foliar and shoot mass for pollarded black locust.     

Demographic variation and habitat specialization of tree species in a diverse tropical forest of Cameroon

Background： Many tree species in tropical forests have distributions tracking local ridge-slope-valley topography. Previous work in a 50-ha plot in Korup National Park, Cameroon, demonstrated that 272 species, or 63% of those tested, were significantly associated with topography. Methods： We used two censuses of 329,000 trees ≥1 cm dbh to examine demographic variation at this site that would account for those observed habitat preferences. We tested two predictions. First, within a given topographic habitat, species specializing on that habitat （＇residents＇） should outperform species that are specialists of other habitats （＇foreigners＇）. Second, across different topographic habitats, species should perform best in the habitat on which they specialize （＇home＇） compared to other habitats （＇away＇）. Species＇ performance was estimated using growth and mortality rates. Results： In hierarchical models with species identity as a random effect, we found no evidence of a demographic advantage to resident species. Indeed, growth rates were most often higher for foreign species. Similarly, comparisons of species on their home vs. away habitats revealed no sign of a performance advantage on the home habitat. Conclusions＂ We reject the hypothesis that species distributions along a ridge-valley catena at Korup are caused by species differences in trees _〉1 cm dbh. Since there must be a demographic cause for habitat specialization, we offer three alternatives. First, the demographic advantage specialists have at home occurs at the reproductive or seedling stage, in sizes smaller than we census in the forest plot. Second, species may have higher performance on their preferred habitat when density is low, but when population builds up, there are negative density-dependent feedbacks that reduce performance. Third, demographic filtering may be produced by extreme environmental conditions that we did not observe during the census interval.     

Patterns of root respiration rates and morphological traits in 13 tree species in a tropical forest

The root systems of forest trees are composed of different diameters and heterogeneous physiological traits. However, the pattern of root respiration rates from finer and coarser roots across various tropical species remains unknown. To clarify how respiration is related to the morphological traits of roots, we evaluated specific root respiration and its relationships to mean root diameter (D) of various diameter and root tissue density (RTD; root mass per unit root volume; gcm(-3)) and specific root length (SRL; root length per unit root mass; mg(-1)) of the fine roots among and within 14 trees of 13 species from a primary tropical rainforest in the Pasoh Forest Reserve in Peninsular Malaysia. Coarse root (2-269mm) respiration rates increased with decreasing D, resulting in significant relationships between root respiration and diameter across species. A model based on a radial gradient of respiration rates of coarse roots simulated the exponential decrease in respiration with diameter. The respiration rate of fine roots (<2mm) was much higher and more variable than those of larger diameter roots. For fine roots, the mean respiration rates for each species increased with decreasing D. The respiration rates of fine roots declined markedly with increasing RTD and increased with increasing SRL, which explained a significant portion of the variation in the respiration among the 14 trees from 13 species examined. Our results indicate that coarse root respiration in tree species follows a basic relationship with D across species and that most of the variation in fine root respiration among species is explained by D, RTD and SRL. We found that the relationship between root respiration and morphological traits provides a quantitative basis for separating fine roots from coarse roots and that the pattern holds across different species.     

Combining canopy height and tree species map information for large-scale timber volume estimations under strong heterogeneity of auxiliary data and variable sample plot sizes

A timber volume regression model applicable to the state and communal forest area of the federal German state of Rhineland-Palatinate is identified using a combination of airborne laser scanning (ALS)-derived metrics and information from a satellite-based tree species classification map available on the federal state level. As is common in many forest inventory datasets, strong heterogeneity in the ALS data due to different acquisition dates and misclassifications in the tree species classification map had noticeable effects on the regression model’s performance. This article specifically addresses techniques that improve the performance of ordinary least square regression models under such restricting conditions. We introduce a calibration technique to neutralize the effect of misclassifications in the tree species variable that originally caused a residual inflation of 0.05 in adjusted \(R^2\). Incorporating the calibrated tree species information improved the model accuracy by up to 0.07 in adjusted \(R^2\) and suggests the use of such information in forthcoming inventories. We also found that including ALS quality information as categorical variables within the regression model considerably mitigates issues with time lags between the ALS and terrestrial data acquisition and ALS quality variations (increase of 0.09 in adjusted \(R^2\)). The model achieved an adjusted \(R^2\) of 0.48 and a cross-validated root-mean-square error (RMSE\(_{\mathrm{cv}}\)) of 46.7% under incorporation of the tree species and ALS quality information and was thus improved by 0.12 in adjusted \(R^2\) (5% in RMSE\(_{\mathrm{cv}}\)) compared to the simple model only containing ALS height metrics (adjusted \(R^2=0.36\), RMSE\(_{\mathrm{cv}}=51.7\)%).     

Topography related spatial distribution of dominant tree species in a tropical seasonal rain forest in China

The degree to which variation in species distribution is predictable from topographic variation is of considerable current interest. In this paper, canonical correspondence analysis (CCA), linear regression and principal coordinates of neighbour matrices (PCNM) models were used to explain the variation in the distributions of the 13 dominant species in a 20-ha tropical rain-forest plot in China. The results showed that: (1) Tree distribution maps show that some species are mainly found in the gullies of the plot, whereas others occur on the slopes. Which indicates topographic variables are important factors for the distribution pattern of species. (2) Both linear regression and CCA results show that convexity and elevation are the most important variables effecting distribution of trees. For saplings, elevation, convexity and aspect explain 15.3%, 9.0% and 10.1% of the total variation of species abundance. For poles, elevation and convexity explain 19.3% and 11.4% respectively. However, only 5.3% of the total variation is explained for adults. (3) The PCNM results showed that topography alone explained 20%, 24% and 5% of the total variation of species abundance for saplings, poles and adults, respectively. Overall evidence for topographic control of the tropical tree distribution is strong, but the explanatory power of topographic variables was a small part of the total of variation.     

Assessing the impacts of species composition, top height and density on individual tree height prediction of quaking aspen in boreal mixedwoods

In this study we developed an individual tree height prediction model for quaking aspen (Populus tremuloides Michx.) grown in boreal mixedwood forests in Alberta using the nonlinear mixed model (NLMM) approach. We examined the impacts of density, species composition, and top height on aspen height predictions. Statistically significant stand level variables were incorporated into the base height–diameter model to increase the predictive ability and accuracy of the model at both the population and subject-specific levels. Our analyses showed that top height and density impacted height growth, but species composition did not. More importantly, we found that the inclusion of additional variables into the base model, despite improving model fitting statistics on the modelling data, did not improve the model's predictive ability and accuracy when cross-validated and when tested on an independent testing data set. Under the NLMM framework the base model performed as well as or better than the expanded models that contained other stand level variables. This has important theoretical and practical implications because, other than for biological reasons, more accurate local tree height predictions for aspen can be achieved simply by using the base height–diameter model fitted with the NLMM approach without the inclusion of other variables.