Irradiance-induced plasticity in the hydraulic properties of saplings of different temperate broad-leaved forest tree species

We assessed the irradiance-related plasticity of hydraulic architecture in saplings of Betula pendula Roth., a pioneer species; Acer pseudoplatanus L., Fraxinus excelsior L. and Quercus robur L., which are post-pioneer light-requiring species; and Quercus petraea Matt. Liebl. and Fagus sylvatica L. Plants were grown in pots in 36%, 16% and 4% of full sunlight. Hydraulic conductance was measured with a high-pressure flow-meter in entire, in situ root systems and in excised shoots. Leaf-specific whole-plant conductance (LSC) increased with irradiance, due, in part, to an effect of irradiance on plant size. In addition, there was a size-independent effect of irradiance on LSC due, in part, to an increase in root hydraulic conductance paralleled by an increase in root biomass scaled to leaf area. Changes in shoot conductivity also contributed to the size-independent plasticity of LSC. Vulnerability to cavitation measured in current-year twigs was much larger in shade-grown plants. Betula pendula had the highest whole-plant, root and shoot conductances and also the greatest vulnerability to cavitation. The other species were similar in LSC, but showed some variation in root conductance scaled to biomass, with Q. robur, Q. petraea and F. sylvatica having the lowest root conductance and susceptibility to cavitation. All species showed a similar irradiance-related plasticity in LSC.     

Leaf-fall phenology along a topography-mediated environmental gradient in a cool–temperate deciduous broad-leaved forest in Japan

Leaf-fall phenology was studied in a cool–temperate deciduous broad-leaved forest in central Japan in relation to the topographic environmental gradients that occur along a short mountain slope. Leaf-fall phenology was described quantitatively using data from leaf litter collected along the slope. In autumn, leaf fall at the study site tended to occur slightly earlier on the upper slope than on the lower slope. This pattern was found at both the stand and the species levels. Our results suggest that leaf-fall phenology may be affected by difference in microclimatic conditions, because environmental conditions are thought to be more severe on the upper slope than on the lower slope. The less intensive methods used in this study, the litter trap method, and Dixon’s model succeeded in quantifying the phenological patterns of leaf fall within stands and within species along the short mountain slope.     

Structural variation in current-year shoots of broad-leaved evergreen tree saplings under forest canopies in warm temperate Japan

Stem length and leaf area of current-year shoots were measured in saplings of eight broad-leaved evergreen tree species growing under a forest canopy. Stem length varied over a range of one to two orders of magnitude within each species. In all species, both the number of leaves and the mean stem length between successive leaves were greater in longer shoots. Mean leaf size and stem length were not correlated in six of eight species, and only weakly positively correlated in the other two species. Thus, total leaf area per stem increased with stem length, but not in direct proportion: leaf area per stem length was smaller in shoots with long stems and larger in shoots with short stems. I conclude that the within-species variation in the leaf-stem balance of current-year shoots is related to variation in shoot functional roles, as has been observed for long and short shoots in many deciduous tree species: shoots with long stems are extension oriented and contribute to the framework of the crown, whereas shoots with short stems serve mainly for leaf display. Among species, large differences were found in the leaf area per stem length ratio. In the species with larger leaf area per stem length ratios, leaves had narrower blades or longer petioles, or both, resulting in a reduction of mutual shading among the leaves on the shoot.     

Evaluation of leaf display of evergreen broadleaved tree species and deciduous tree species in warm temperate conifer plantations

We investigated the sapling leaf display in the shade among trees of various leaf lifespans co-occurring under the canopy of a warm-temperate conifer plantation. We measured leaf-area ratio (aLAR) and morphological traits of saplings of evergreen broadleaved tree species and a deciduous tree species. Although we found large interspecific and intraspecific differences in aLAR even among saplings of similar size in the homogeneous light environment, we did not find a consistent trend in aLAR with leaf lifespan among the species. While deciduous trees annually produced a large leaf area, some evergreen broadleaved trees retained their leaves across years and had aLAR values as high as those of deciduous trees. Among leaf-level, shoot-level, and individual-level morphological traits, aLAR was positively correlated with current-year shoots mass per aboveground biomass in deciduous trees, and with the area of old leaves per aboveground mass in evergreen broadleaved trees. Thus, tree-to-tree variation in the degrees of annual shoot production and the accumulation of old leaves were responsible for the interspecific and intraspecific variations in aLAR.     

Impacts of temperate lianas on tree growth in young deciduous forests

Lianas are often overlooked in temperate ecological studies even though they are important components of forest communities. While lianas have been shown to damage tropical canopy trees and reduce the growth of juvenile trees, the impact of lianas on canopy tree growth in temperate systems is largely unknown. Growth of trees ≥8 cm dbh was examined over a 9-year period within 50-year old post-agricultural secondary forests in the Piedmont region of New Jersey, USA. Five lianas, Celastrus orbiculatus, Lonicera japonica, Parthenocissus quinquefolia, Toxicodendron radicans, and Vitis species, occurred throughout the forest. Total liana basal area, number of stems, and percent cover within host trees were evaluated to assess liana burdens on 606 previously censused trees. These data were related to tree growth to assess liana impacts. Forest trees were separated based on their dominance in the canopy to determine whether lianas had the potential to influence forest composition. In general, lianas in the forests were fairly abundant, with 68% of the trees having at least one liana present. On average, each tree supported 9.7 cm² of liana basal area and 23% of the canopy was covered by lianas. Most of the variation in tree growth was related to the dominance of trees within the canopy, with canopy dominant and co-dominant trees growing 2.5× more than suppressed trees. Liana basal area and number of lianas stems were not related to tree growth, but liana canopy cover decreased tree growth. However, not all trees were equally affected as canopy cover of lianas only reduced growth in dominant and co-dominant trees. Lianas were most influential on host tree growth in unsuppressed trees when occupying a majority of the canopy, only a minority of forest trees. This suppression was not related to differential liana colonization of canopy trees as all canopy classes supported equivalent liana burdens. Though lianas impacted only a minority of the trees in this system, some liana species, C. orbiculatus and Vitis spp., are still increasing and may pose future risks to forest growth and development.     

Physiological basis of seasonal trend in leaf photosynthesis of five evergreen broad-leaved species in a temperate deciduous forest

The physiological basis of photosynthesis during winter was investigated in saplings of five evergreen broad-leaved species (Camellia japonica L., Cleyera japonica Thunb., Photinia glabra (Thunb.) Maxim., Castanopsis cuspidata (Thunb.) Schottky and Quercus glauca Thunb.) co-occurring under deciduous canopy trees in a temperate forest. We focused on temperature dependence of photosynthetic rate and capacity as important physiological parameters that determine light-saturated rates of net photosynthesis at low temperatures during winter. Under controlled temperature conditions, maximum rates of ribulose bisphosphate carboxylation and electron transport (Vcmax) and Jmax, respectively) increased exponentially with increasing leaf temperature. The temperature dependence of photosynthetic rate did not differ among species. In the field, photosynthetic capacity, determined as Vcmax and Jmax at a common temperature of 25 degrees C (Vcmax(25) and Jmax(25)), increased until autumn and then decreased in species-specific patterns. Values of Vcmax(25) and Jmax(25) differed among species during winter. There was a positive correlation of Vcmax(25) with area-based nitrogen concentration among leaves during winter in Camellia and Photinia. Interspecific differences in Vcmax(25) were responsible for interspecific differences in light-saturated rates of net photosynthesis during winter.     

Productivity of temperate broad-leaved forest stands differing in tree species diversity

? Understanding the effects of tree species diversity on biomass and production of forests is fundamental for carbon sequestration strategies, particularly in the perspective of the current climate change. However, the diversity-productivity relationship in old-growth forests is not well understood.

? We quantified biomass and above-ground production in nine forest stands with increasing tree species diversity from monocultures of beech to stands consisting of up to five deciduous tree species (Fagus sylvatica, Fraxinus excelsior, Tilia spp., Carpinus betulus, Acer spp.) to examine (a) if mixed stands are more productive than monospecific stands, (b) how tree species differ in the productivity of stem wood, leaves and fruits, and (c) if beech productivity increases with tree diversity due to lower intraspecific competition and complementary resource use.

? Total above-ground biomass and wood production decreased with increasing tree species diversity. In Fagus and Fraxinus, the basal area-related wood productivity exceeded those of the co-occurring tree species, while Tilia had the highest leaf productivity. Fagus trees showed no elevated production per basal area in the mixed stands.

? We found no evidence of complementary resource use associated with biomass production. We conclude that above-ground productivity of old-growth temperate deciduous forests depend more on tree species-specific traits than on tree diversity itself.

     

Structure and composition of the seed bank in monospecific and tree species-rich temperate broad-leaved forests

How tree species diversity affects ecosystem functioning is a topic of intensive research. This study compares monospecific and species-rich broad-leaved forests under similar bedrock and climate conditions for the size and composition of their seed bank. We tested the hypotheses that (i) the actual herb-layer vegetation has an only weak influence on the composition of the seed bank, (ii) the species diversity of the seed bank increases with tree-layer diversity, and (iii) tree species forming a more persistent litter layer reduce the number of germinating seeds. The number of seeds and their species composition were investigated in soil cores taken from three soil depths (0–5, 5–10 and 10–20 cm; n = 4, with each 6 sub-samples) in 9 study plots differing in tree species diversity (3 monospecific Fagus plots, 3 plots with Fagus, Fraxinus and Tilia, and 3 plots with Fagus, Fraxinus, Tilia, Carpinus and Acer). Tree species diversity had a much stronger influence on the size and composition of the seed bank than herb-layer diversity or composition, the latter revealing only a low similarity to the corresponding seed bank. The number and species diversity of emerging seedlings decreased significantly with the amount of acidifying Fagus litter, but increased with litter mass of Tilia and other trees with nutrient-rich, rapidly decomposing litter. We conclude that tree species diversity does not influence the seed bank through effects on herb-layer composition, but mostly through differential disturbance histories of the stands and litter quality effects on germination and soil chemistry. From the contrasting effects of Fagus and Tilia leaf litter, it appears that effects of tree species identity are more relevant than influences of tree species diversity itself.     

蒙古栎林在温带落叶阔叶林群落分级中的地位

为清楚阐明蒙古栎林的群落分级及其在温带落叶阔叶林群落分级中的地位，我们收集了东北亚201个蒙古栎林和日本的153个栎林的资料．将其综合于一个简表中．并应用Braun—Blanquet方法对这些森林植被进行了比较。结果表明：在温带落叶阔叶林的群落分级系统中。蒙古栎林与水青冈纲占据着同等重要的地位，它们共同构成温带落叶阔叶林纲群(Class-group)。不同生境下的蒙古栎的20个群集或群落被归纳成为2个目、3个群团和3个亚群团。     

Allometric differences between current-year shoots and large branches of deciduous broad-leaved tree species

Current-year shoots are mostly made of primary tissues, whereas first-order branches comprise mainly secondary tissues. Differences in tissue composition of these units reflect differences in functional design. We compared the allometry of current-year shoots and first-order branches in eight deciduous broad-leaved tree species and examined the functional differences underlying the design of current-year shoots and first-order branches. Allometric relationships of first-order branches tended to be compatible with predictions of the pipe model and elastic similarity model. That is, allometric constants of the relationships between leaf mass and stem diameter at the branch base and between stem diameter and stem mass were 2.0 and 0.33-0.38, respectively, indicating that the functional regulation of stem form of first-order branches can be predicted by the two models. However, allometric relationships of current-year shoots were not compatible with the predictions of the pipe and elastic similarity models. Thus, the allometric constant of the relationship between leaf mass and stem diameter at the base of current-year shoots was larger than 2.0, and the allometric constant of the relationship between stem length and stem diameter of shoots was larger than 1.0 in all species examined. However, current-year shoots had an allometric constant of leaf mass against stem length that was less than 1.0, suggesting a functional demand on shoot design to reduce self-shading. Also, allometric constants of stem length against stem diameter at the shoot base were larger in monopodial species than in sympodial species, whereas allometric constants of leaf mass per shoot against stem length were smaller in monopodial species than in sympodial species. We propose that the allometries of current-year shoots reflect their function as disposable units for temporary leaf arrangement.     

Demographic and height growth response of native broad-leaved deciduous tree saplings to overhead canopy release in a coastal Pinus thunbergii forest in Hokkaido, northern Japan

Prevention efforts to combat pine wilt disease must be instituted to avoid destruction of coastal forests in Hokkaido, northern Japan. We examine an alternative method involving the conversion of Pinus thunbergii forests to native broadleaf species through advancing their growth, but we have little knowledge about the response of advanced growth to overhead canopy release. To demonstrate the demographic and height growth response of advanced growth to the light change in a coastal Pinus thunbergii forest, canopy trees were removed and the sapling populations were monitored for 6?years. Inhibition in height growth caused by light increase was observed in Cerasus sargentii and Kalopanax septemlobus saplings. Especially in Kalopanax septemlobus, a remarkable decline was observed in the saplings growing in the darker positions which experienced light change. Therefore, large light changes should be carefully avoided for this species especially in the darker positions. In contrast, relative height growth rates of Quercus crispula and, especially, Quercus dentata increased with increasing light, and saplings tended to die in the darker positions. No dead trees of Sorbus commixta were observed during the study period. The saplings also showed good height growth even when under closed canopy. This species seems to be adaptive to a dark environment. Since Quercus dentata and Kalopanax septemlobus are the major components of the natural coastal forests in Hokkaido, gradual canopy release is available to foster advanced growth in coastal Pinus thunbergii forest, in accordance with the concept of density control in coastal forests.     

Environmental control of daily stem growth patterns in five temperate broad-leaved tree species

Tree ring analysis investigates growth processes at time horizons of several weeks to millennia, but lacks the detail of short-term fluctuation in cambial activity. This study used electronic high-precision dendrometry for analyzing the environmental factors controlling stem diameter variation and radial growth in daily resolution in five co-existing temperate broad-leaved tree species (genera Fraxinus, Acer, Carpinus, Tilia and Fagus) with different growth and survival strategies. Daily stem radius change (SRC(d)) was primarily influenced by the atmospheric demand for water vapor (expressed either as vapor pressure deficit (D) or relative air humidity (RH)) while rainfall, soil matrix potential, temperature and radiation were only secondary factors. SRC(d) increased linearly with increasing RH and decreasing D in all species. The positive effect of a low atmospheric water vapor demand on SRC(d) was largest in June during the period of maximal radial growth rate and persisted when observation windows of 7 or 21 days instead of 1 day were used. We found a high synchronicity in the day-to-day growth rate fluctuation among the species with increment peaks corresponding to air humidity maxima, even though the mean daily radial growth rate differed fivefold among the species. The five -species also differed in the positive slope of the growth/RH relationship with the steepest increase found in Fraxinus and the?lowest in Fagus. We explain the strong positive effect of high RH and low D on radial stem increment by lowered transpiration which reduces negative pressure in the conducting system and increases turgor in the stem cambium cells, thereby favoring cell division and expansion. The results suggest that mechanistic models of tree growth need to consider the atmospheric water status in addition to the known controlling environmental factors: temperature, soil moisture and precipitation. The results further have implications for sensitivity analyses of tree growth to climatic changes.     

Dark respiration rate increases with plant size in saplings of three temperate tree species despite decreasing tissue nitrogen and nonstructural carbohydrates

In shaded environments, minimizing dark respiration during growth could be an important aspect of maintaining a positive whole-plant net carbon balance. Changes with plant size in both biomass distribution to different tissue types and mass-specific respiration rates (R(d)) of those tissues would have an impact on whole-plant respiration. In this paper, we evaluated size-related variation in R(d), biomass distribution, and nitrogen (N) and total nonstructural carbohydrate (TNC) concentrations of leaves, stems and roots of three cold-temperate tree species (Abies balsamea (L.) Mill, Acer rubrum L. and Pinus strobus L.) in a forest understory. We sampled individuals varying in age (6 to 24 years old) and in size (from 2 to 500 g dry mass), and growing across a range of irradiances (from 1 to 13% of full sun) in northern Minnesota, USA. Within each species, we found small changes in R(d), N and TNC when comparing plants growing across this range of light availability. Consistent with our hypotheses, as plants grew larger, whole-plant N and TNC concentrations in all species declined as a result of a combination of changes in tissue N and shifts in biomass distribution patterns. However, contrary to our hypotheses, whole-plant and tissue R(d) increased with plant size in the three species.     

Ontogenetic changes in leaf phenology of Ulmus davidiana var. japonica, a deciduous broad-leaved tree

To determine how plants control leaf phenology to maximize annual carbon gain, I examined ontogenetic changes in leaf phenology of Japanese elm, Ulmus davidiana var. japonica Nakai plants of different ages growing in contrasting light environments. Leaf emergence occurred earlier in 1- and 2-year-old seedlings than in current-year seedlings. Although leaf emergence was not affected by light conditions at the sites, it was influenced by plant height. The delay in leaf emergence increased with increasing plant height. These traits indicate that seedlings that received the least light during the summer intercepted light for a long period during the spring; however, the advantage of earlier leaf emergence decreased with increasing plant height. At each site, 1-year-old seedlings had a longer duration of leaf emergence than adults, because of a longer period of favorable light conditions even in the forest understory. Duration of leaf emergence, leaf duration and leaf longevity were usually longer in sun than in shade for both seedlings and adults; however, flexibility in the response to light was greater in seedlings than in adults. The plastic response in leaf phenology during the juvenile stages may contribute to the optimization of light acquisition in habitats with differing light conditions, thereby enhancing seedling survival.     

Decomposition of leaf litter of four native broad-leaved tree species in south China

Leaf litter decomposition of Castanopsis fissa, Cinnamomum camphora, Michelia macclurei and Mytilaria laosensis in mixed broad-leaved plantation and pine plantation was studied by the litterbag method for 1 year. Leaf litter decomposition rates of the four species were highest in Cinnamomum camphora, followed by Mytilaria laosensis, Michelia macclurei, and Castanopsis fissa. The decomposition rates of all four species were higher in the mixed than in pine plantation. The decomposition processes of all species followed Olson’s exponential model. The decomposition coefficients (k) of all species were also higher in the mixed plantation and had the same order as the decomposition rates. The nitrogen contents of leaf litter of the different species studied increased initially and then decreased with time. Net release of N only occurred in pine plantation. Potassium contents appeared to decrease first but later increase, and net release was only found in mixed plantation. Calcium, magnesium and boron all showed similar pattern of initial increase followed by later decrease. They all had net release in both mixed and pine plantations. The release of phosphorus varied greatly between species and showed no clear trend.     

Direct effect of elevated CO(2) on nocturnal in situ leaf respiration in nine temperate deciduous tree species is small

Direct (i.e., short-term) effects of elevated CO(2) on nocturnal in situ leaf respiration rate were measured in nine deciduous tree species (seven genera) in 20 3.5-4.0-h experiments. During the experiments, CO(2) concentration was alternated between 400 and 800 ppm (approximately 40 and 80 Pa of CO(2)). Data analysis accounted for effects on respiration rate of the normal decline in temperature with time after sunset. The median response to a 40-Pa increase in CO(2) was a 1.5% decrease in respiration rate, with responses ranging from a 5.6% inhibition to a 0.4% stimulation. Direct effects of elevated CO(2) on respiration were similar among the species. Thus, the response of nocturnal leaf respiration rate to a short-term CO(2) increase was small, and of little practical importance to the accuracy of measurements of respiration involving similar changes in CO(2) concentration during measurement. These direct respiratory responses of leaves to elevated CO(2) would translate into only slight, if any, effects on the carbon balance of temperate deciduous forests in a future atmosphere containing as much as 80 Pa CO(2).     

Seasonal fluctuations and temperature dependence of leaf gas exchange parameters of co-occurring evergreen and deciduous trees in a temperate broad-leaved forest

Seasonal fluctuations in leaf gas exchange parameters were investigated in three evergreen (Quercus glauca Thunb., Cinnamomum camphora Sieb. and Castanopsis cuspidata Schottky) and one deciduous (Quercus serrata Thunb.) co-occurring, dominant tree species in a temperate broad-leaved forest. Dark respiration rate (Rn), maximum carboxylation rate (Vcmax) and stomatal coefficient (m), the ratio of stomatal conductance to net assimilation rate after adjustment to the vapor pressure deficit and internal carbon dioxide (CO2) concentration, were derived inversely from instantaneous field gas exchange data (one-point method). The normalized values of Rn and Vcmax at the reference temperature of 25 degrees C (Rn25, Vcmax25) and their temperature dependencies (Delta Ha(Rn), Delta Ha(Vcmax)) were analyzed. Parameter Vcmax25 ranged from 24.0-40.3 micromol m(-2) s(-1) and Delta Ha(Vcmax) ranged from 29.1- 67.0 kJ mol(-1). Parameter Rn25 ranged from 0.6-1.4 micromol m(-2) s(-1) and Delta Ha(Rn) ranged from 47.4-95.4 kJ mol(-1). The stomatal coefficient ranged from 7.2-8.2. For the three evergreen trees, a single set of Vcmax25 and Rn25 parameters and temperature dependence curves produced satisfactory estimates of carbon uptake throughout the year, except during the period of simultaneous leaf fall and leaf expansion, which occurs in April and May. In the deciduous oak, declines in Vcmax25 were observed after summer, along with changes in Vcmax25 and Rn25 during the leaf expansion period. In all species, variation in m during periods of leaf expansion and drought should be considered in modeling studies. We conclude that the changes in normalized gas exchange parameters during periods of leaf expansion and drought need to be considered when modeling carbon uptake of evergreen broad-leaved species.     

Hydraulic properties and embolism in small-diameter roots of five temperate broad-leaved tree species with contrasting drought tolerance

? Context

It has been estimated that about half of a plant??s total hydraulic resistance is located belowground, but it is not well known how temperate tree species differ in root hydraulic properties and how these traits vary with the species?? drought tolerance.

? Aims

We examined root anatomical and hydraulic traits in five broad-leaved tree species with different drought tolerance, analyzed the relation between root anatomy and hydraulic conductivity and root embolism, and investigated the relation of these traits to the species?? drought tolerance.

? Methods

In small-diameter roots (2?C6?mm), we measured vessel diameters and vessel density, specific hydraulic conductivity, and the percental loss of conductivity (??native?? embolism) during summer in a mixed forest.

? Results

Specific conductivity was positively related to vessel diameter but not to vessel density. Drought-tolerant Fraxinus showed the smallest mean vessel diameters and drought-sensitive Fagus the largest. Specific conductivity was highly variable among different similar-sized roots of the same species with a few roots apparently functioning as ??high-conductivity roots??.

? Conclusion

The results show that coexisting tree species can differ largely in root hydraulic traits with more drought-sensitive trees apparently having larger mean vessel diameters in their roots than tolerant species. However, this difference was not related to the observed root conductivity losses due to embolism.