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Variations in transpiration rate and leaf cell turgor maintenance in saplings of deciduous broad-leaved tree species common in cool temperate forests in Japan.

To clarify mechanisms underlying variation in transpiration rate among deciduous broad-leaved tree species, we measured diurnal changes in stomatal conductance (gs) and leaf water potential, and calculated the maximum transpiration rate (Emax), leaf-specific hydraulic conductance (K(s-l)) and difference between the soil water potential and the daily minimum leaf water potential (Psis - Psi(l,min)). Pressure-volume (P-V) measurements were made on leaves. Saplings of eight broad-leaved tree species that are common in Japanese cool temperate forests were studied. Maximum transpiration rate varied significantly among species. There was a statistically significant difference in Psis - Psi(l,min), but not in K(s-l). Species with large Emax also had large Psis - Psi(l,min) and gs. The results of the P-V analyses showed that species with a large Psis - Psi(l,min) maintained turgor even at low leaf water potentials. The similar daily minimum leaf pressure potentials (Psip) across all eight species indicate that Psip values below this minimum are critical. Based on these results, we suggest that the leaf cell capacity for turgor maintenance strongly affects Psis - Psi(l,min) and consequently Emax via stomatal regulation.

Betula↗

Interactive effects of temperature and precipitation on soil respiration in a temperate maritime pine forest.

Soil respiration (SR) was monitored periodically throughout 2001 in a Scots pine (Pinus sylvestris L.) stand located in the Belgian Campine region. As expected for a temperate maritime forest, temperature was the dominant control over SR during most of the year. However, during late spring and summer, when soil water content (SWC) was limiting, SR was insensitive to temperature (Q(10) = 1.24). We observed that during prolonged rain-free periods, when SWC was less than 15% (v/v), SR decreased dramatically (up to 50%) and SWC took over control of SR. During such drought periods, however, rain events sometimes stimulated SR and restored temperature control over SR, even though SWC in the mineral soil was low. We hypothesize that restoration of temperature control occurred only when rain events adequately rewetted the uppermost soil layers, where most of the respiratory activity occurred. To quantify the rewetting capacity of rain events, an index (I(w)) was designed that incorporated rainfall intensity, time elapsed since the last rain event, and atmospheric vapor pressure deficit (a proxy for evaporative water losses). To simulate SR fluxes, a model was developed that included the effects of soil temperature and, under drought and non-rewetting conditions (I(w) and SWC < threshold), an SWC response function. The model explained 95% of the temporal variability in SR observed during summer, whereas the temperature function alone explained only 73% of this variability. Our results revealed that, in addition to temperature and SWC, rain plays a role in determining the total amount of carbon released from soils, even in a maritime climate.

Cell Respiration↗

In situ measurement of water absorption by fine roots of three temperate trees: species differences and differential activity of superficial and deep roots.

The spatial heterogeneity of water uptake by fine roots under field conditions was analyzed in situ with miniature sap flow gauges in a mature beech-oak-spruce mixed stand. Sap flow rate (J), sap flow density (Jd), and root surface-area-specific flow rate (uptake rate, Js) were measured for eight to 10 small-diameter roots (3-4 mm) per species in the organic layer (superficial roots) and in the mineral soil (30-80 cm, deep roots) during four months in summer 1999. We calculated Js by relating J to the surface area of the section of the fine root system distal to the position of the gauge on the root. When measured synchronously, roots of the three species did not differ significantly in mean Js, although oak roots tended to have lower rates. However, Jd decreased in the sequence spruce > beech > oak in most measurement periods. Microscopic investigation revealed differences in fine root anatomy that may partly explain the species differences in Jd and Js. Oak fine roots had a thicker periderm than beech and spruce roots of similar diameter and spruce roots had fewer fine branch rootlets than the other species. Synchronously recorded Jd and Js of nearby roots of the same tree species showed large differences in flow with coefficients of variation from 25 to 150% that could not be explained by patchy distribution of soil water. We hypothesize that the main cause of the large spatial heterogeneity in root water uptake is associated with differences between individual roots in morphology and ultrastructure of the root cortex that affect root radial and root-soil interface conductivities. The high intraspecific variation in Js may mask species differences in root water uptake. Superficial roots of all species typically had about five times higher Jd than deep roots of the same species. However, Js values were similar for superficial and deep roots in beech and spruce because small diameter roots of both species were more branched in the organic layer than in mineral soil. In oak, deep roots had lower Js (maximum of 100 g m(-2) day(-1)) than superficial roots (about 1000 g m(-2) day(-1)). We conclude that temperate tree species in mixed stands have different water uptake capacities. Water flow in the rhizosphere of forests appears to be a highly heterogeneous process that is influenced by both tree species and differences in uptake rates of individual roots within a species.

Fagus↗

Water storage capacitance and xylem tension in isolated branches of temperate and tropical trees.

Trees of tropical semi-deciduous forests range from "drought-avoiding" stem-succulent species with low-density wood (< 0.5 g cm(-3)), which maintain high stem water potentials (psi(STEM) > -0.7 MPa) throughout the year, to "drought-tolerant" deciduous hardwood species (wood density > 0.75 g cm(-3)), which dehydrate strongly during seasonal drought (psi(STEM) < -6 MPa). In stem-succulent and other drought-avoiding species, xylem vessels are surrounded by extensive parenchyma providing intracellular water storage, whereas in deciduous species stem water storage is mainly extracellular. Thirteen tropical and two temperate tree species, representing different functional types, were studied. The contribution of stem water storage to these species' water use during water stress was determined by time-series analysis of dehydration and rehydration of excised leaf-bearing branches of these trees. During dehydration, stem water potential slowly declined 1-2 MPa in drought-avoiding species, but in deciduous species it rapidly fell 4-5 MPa, suggesting that water storage capacitance was related to xylem anatomy. After immersion of dehydrated, leafless branches in water, the decline in xylem tension and rate of water uptake during rehydration were linearly related, as predicted by application of Ohm's law to water flux. The decline of xylem tension during rehydration was biphasic, with a phase of rapid water uptake into extracellular spaces being followed by a prolonged phase of slow water uptake into living cells. The rate of water uptake during rehydration and the minima of leaf water potential observed in the field during the dry season were highly correlated with water storage capacitance, indicating that wood anatomy is a major determinant of drought adaptation.

Climate↗

Species differences in timing of leaf fall and foliage chemistry modify nutrient resorption efficiency in deciduous temperate forest stands.

Extensive variation in fractional resorption of mineral elements from plant leaves is still not fully understood. In multi-species forest stands, species leaf fall phenology and leaf constitution may significantly modify the timing of nutrient return to the soil and overall plant nutrient loss. We studied leaf fall and nutrient loss kinetics, and leaf composition in three natural, temperate, deciduous broadleaf forest stands to determine the role of timing of leaf abscission and nutrient immobilization in cell walls on nutrient resorption efficiency of senescing leaves. Nitrogen (N), phosphorus and potassium contents decreased continuously in attached leaves after peak physiological activity during mid-season. Changes in nutrient contents of attached leaves were paralleled by decreases in nutrient contents in freshly fallen leaf litter. In different species and for different nutrients, resorption of nutrients from senescing leaves proceeded with different kinetics. The maximum nutrient resorption efficiency (the fraction of specific nutrient resorbed from the leaves at the end of leaf fall) did not depend on the mid-seasonal nutrient concentration. Species with earlier leaf fall resorbed leaf nutrients at a faster rate, partly compensating for the earlier leaf fall. Nevertheless, the litter-mass weighted mean nutrient contents in leaf litter were still larger in species with earlier leaf fall, demonstrating an inherent trade-off between early leaf fall and efficient nutrient resorption. This trade-off was most important for N. Losses of the non-mobile nutrients calcium and magnesium were unaffected by the timing of leaf fall. There was large variation in the maximum N resorption efficiency among species. Correlations among leaf chemical variables suggested that the maximum N resorption efficiency decreased with the increasing fraction of cell walls in the leaves, possibly due to a greater fraction of N occluded in cell wall matrix. We conclude that species leaf fall phenology and leaf chemistry modify the timing and quantities of plant nutrient losses, and that more diverse forest stands supporting a spectrum of species with different phenologies and leaf types produce litter with more variable chemical characteristics than monotypic stands.

Climate↗

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.

Algorithms↗

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.

Abies↗

Models of the spring phenology of boreal and temperate trees: Is there something missing?

According to prevailing theory, air temperature is the main environmental factor regulating the timing of bud burst of boreal and temperate trees. Air temperature has a dual role in this regulation. First, after the cessation of growth in autumn, prolonged exposure to chilling causes rest completion, i.e., removes the physiological growth-arresting conditions inside the bud. After rest completion, prolonged exposure to warm conditions causes ontogenetic development leading to bud burst or flowering. During the past three decades, several simulation models based on chilling and forcing have been developed and tested. In recent modeling studies of the timing of bud burst in mature trees, the simpler thermal-time models that assume forcing starts on a fixed date in the spring have outperformed the chilling-forcing models. We hypothesize that this discrepancy may be due to some element missing from the chilling-forcing models. We tested two new model formulations by introducing reversing, temperature-driven elements that precede forcing and by fitting the models to seven historical time series of data of flowering and leaf bud burst of common boreal tree species. In these tests, both of the new models were generally more accurate in predicting the timing of bud burst than a classical chilling-forcing model, but less accurate than the simple thermal-time model. We therefore conclude that besides chilling, other environmental factors are involved in the regulation of the timing of bud burst. Further work is needed to determine if the regulatory factors derive from air temperature or from some other environmental condition such as changes in light conditions, like day length or night length.

Alnus↗

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.

Climate↗

Response of five temperate deciduous tree species to water stress.

Gas exchange, tissue water relations, and leaf/root dry weight ratios were compared among young, container-grown plants of five temperate-zone, deciduous tree species (Acer negundo L., Betula papyrifera Marsh, Malus baccata Borkh, Robinia pseudoacacia L., and Ulmus parvifolia Jacq.) under well-watered and water-stressed conditions. There was a small decrease (mean reduction of 0.22 MPa across species) in the water potential at which turgor was lost (Psi(tlp)) in response to water stress. The Psi(tlp) for water-stressed plants was -1.18, -1.34, -1.61, -1.70, and -2.12 MPa for B. papyrifera, A. negundo, U. parvifolia, R. pseudoacacia, and M. baccata, respectively. Variation in Psi(tlp) resulted primarily from differences in tissue osmotic potential and not tissue elasticity. Rates of net photosynthesis declined in response to water stress. However, despite differences in Psi(tlp), there were no differences in net photosynthesis among water-stressed plants under the conditions of water stress imposed. In A. negundo and M. baccata, water use efficiency (net photosynthesis/transpiration) increased significantly in response to water stress. Comparisons among water-stressed plants showed that water use efficiency for M. baccata was greater than for B. papyrifera or U. parvifolia. There were no significant differences in water use efficiency among B. papyrifera, U. parvifolia, A. negundo, and R. pseudoacacia. Under water-stressed conditions, leaf/root dry weight ratios (an index of transpiration to absorptive capacity) ranged from 0.77 in R. pseudoacacia to 1.05 in B. papyrifera.

Journal Article↗

Considerations in modeling change in temperate forest nitrogen cycles.

The general features of nitrogen (N) cycles in temperate forests and the important processes to consider when modeling change in these cycles include atmospheric inputs, N fixation, litter (root and aboveground) transfers and decomposition, soil processes, N uptake and effects on productivity and litter quality, and N outputs. Nitrogen cycling is closely linked with the carbon (C) and water cycles. Thus models of N cycling must include aspects of these other cycles. Although much is known about individual processes, development of a generic model of forest N cycling is not possible at present because the links and interactions among the individual processes are not well understood. The weakest links with respect to the N cycle are: quantification of atmospheric (especially dry) deposition rates in polluted environments, controls on C and N allocation in vegetation, controls on N turnover in fine roots, controls on decomposition of the older components of soil organic matter, and feedbacks among N availability, litter "quality" and subsequent N mineralization rates. To examine possible effects of long-term change in climate or atmospheric chemistry on the storage of C and other elements in forest ecosystems, we need to model in detail the effects of these factors on complex soil processes such as organic matter decomposition. Some promising models have been developed, but they need to be validated across a range of forest types before they can be used with confidence for long-term prediction. Mean annual leaf litter N concentration offers potential as a simple index of annual N uptake in forests.

Journal Article↗

Shrimps remove ectoparasites from fishes in temperate waters.

We have found that two very common species of North Atlantic shallow water shrimp, Palaemon adspersus and Palaemon elegans, remove and feed on ectoparasites on plaice (Pleuronectes platessa L.). The relationship could be mutualistic, as we did not observe any attempts by the fishes to feed on the shrimps. The ectoparasites removed included monogenean worms (Gyrodactylus sp.) and sea lice (Lepeophtheirus pectoralis). An experiment showed that there were 65% more Gyrodactylus parasites on the fishes that had been apart from compared with those that had been together with shrimps for 48h. Shrimps on coral reefs are known for cleaning fishes, but that shrimps in temperate waters show parasite-cleaning behaviour is, to our knowledge, a new observation.

Animals↗

Sex and segregation in temperate bats.

Many temperate insectivorous bats show marked sexual segregation during the summer, but in spectacular, pre-hibernation swarming, gather at caves to mate. In many species, sexual segregation is probably due to a gradient in aerial insect availability that confines females to lower elevations, where high reproductive costs are met by an abundant and reliable food supply. In the hawking and trawling Myotis daubentonii, we show that alongside inter-sexual segregation, there is intra-male segregation and suggest that this results from the exclusion of most males from high-quality habitat. These apparently excluded males suffer reduced foraging efficiency and mating success relative to males that roost with the females in summer. Changes in resources and behaviour at the end of the summer lead to a change in strategy that gives all males a chance to mate during swarming, but this does not overcome the paternity advantage to males that spend the summer with the females.

Animals↗

Coherent assembly of phytoplankton communities in diverse temperate ocean ecosystems.

The annual cycle of phytoplankton cell abundance is coherent across diverse ecosystems in the temperate North Atlantic Ocean. In Bedford Basin, on the Scotian Shelf and in the Labrador Sea, the numerical abundance of phytoplankton is low in spring and high in autumn, thus in phase with the temperature cycle. Temperature aligns abundance on a common basis, effectively adjusting apparent cell discrepancies in waters that are colder or warmer than the regional norm. As an example of holistic simplicity arising from underlying complexity, the variance in a community variable (total abundance) is explained by a single predictor (temperature) to the extent of 75% in the marginal seas. In the estuarine basin, weekly averages of phytoplankton and temperature computed from a 13 year time-series yield a predictive relationship with 91% explained variance. Temperature-directed assembly of individual phytoplankton cells to form communities is statistically robust, consistent with observed biomass changes, amenable to theoretical analysis, and a sentinel for long-term change. Since cell abundance is a community property in the same units for all marine microbes at any trophic level and at any phylogenetic position, it promises to integrate biological oceanography into general ecology and evolution.

Atlantic Ocean↗

Modelling dispersal of a temperate insect in a changing climate.

We construct a novel individual-based random-walk model to assess how predicted global climate change might affect the dispersal rates of a temperate insect. Using a novel approach we obtained accurate field measurements of daily movements for individuals over time to parameterize our model. Males were found to move significantly further on average than females. Significant variation in movement was evident among individuals; the most dispersive individuals moved up to five (females) and seven (males) times as far on average as the least dispersive individuals. Mean relative daily movement of both males and females were exponentially related to maximum daily temperature recorded within the grass sward. Variability, both within and among individuals, in relative daily movement was incorporated into the model using gamma probability distributions. Resultant dispersal functions for seasonal movement are predicted to be highly leptokurtic, which agrees well with observations from the field. Predictions of the model suggest that for populations at the polewards edge of the current range an increase of 3-5 degrees C in daily maximum temperature may increase the proportion of long-distance dispersers (those characterized as comprising the top 0.1% of furthest dispersing individuals under local conditions experienced during the 1963-1990 period) by up to 70%.

Animals↗

Economic, biological and policy constraints on the adoption of carbon farming in temperate regions.

In this paper the issues surrounding the potential role for agriculture in temperate climates in the mitigation of greenhouse gases are examined, with a particular focus on the constraints and limitations on the adoption of practices contributing to carbon sequestration. Other land uses have come under close scrutiny for their potential to act as carbon sinks and it is likely that soil sequestration may become a legitimate part of the 'Land-use, land-use change and forestry' mechanism. However, for this to occur, further developments in our understanding of the biological processes involved in soil-carbon sequestration are required.

Agriculture↗

Patterns in the assembly of temperate forests around the Northern Hemisphere.

Recent studies of Northern Hemisphere biogeography have highlighted potentially significant differences between disjunction patterns in plants versus animals. To assess such differences, we compiled a larger sample of relevant plant phylogenies from which disjunction patterns, ancestral areas and directions of movement could be inferred. We considered 66 plant clades with species variously endemic today to eastern Asia (EA), Europe (including southwestern Asia), eastern North America (ENA), and/or western North America (WNA). Within these clades we focused on 100 disjunctions among these major areas, for 33 of which absolute divergence times have also been inferred. Our analyses uphold the view that disjunctions between EA and ENA are exceptionally common in plants, apparently more so than in animals. Compared with animals, we find few disjunctions between EA and WNA, consistent with increased extinction in WNA or failure of some groups to colonize that region. Taken at face value, our data also support the view that many temperate forest plant groups originated and diversified within EA, followed by movement out of Asia at different times, but mostly during the last 30 Myr. This favours Beringia over a North Atlantic land bridge as the primary path between the Old World and the New World. Additional studies are needed, especially to evaluate the impacts of differential extinction on these patterns, to more confidently establish divergence times, and to assess the statistical significance of these findings. Fortunately, many more plant groups show relevant disjunction patterns and could soon be added to such analyses.

Asia↗

Shewanella olleyana sp. nov., a marine species isolated from a temperate estuary which produces high levels of polyunsaturated fatty acids.

Two polyunsaturated fatty acid (PUFA) producing strains (ACEM 6 and ACEM 9(T)) isolated from a temperate, humic-rich river estuary in Tasmania, Australia, were found to be members of the genus Shewanella. These strains were able to utilize humic compounds (tannic acid) and derivatives (2,6-anthraquinone disulfonate) as sole carbon sources and as electron acceptors for anaerobic respiration. The major fatty acids were typical of the genus Shewanella; however, PUFAs mostly made up of eicosapentaenoic acid were produced at high levels (10.2-23.6% of total fatty acids) and at relatively high incubation temperatures (10.2% at 24 degrees C). Sequence analysis indicated that ACEM 6 and ACEM 9(T) had identical 16S rDNA sequences and were most closely related to Shewanella japonica (sequence similarity 97.1%). DNA hybridization and phenotypic characteristics confirmed that the isolates constituted a novel species of the genus Shewanella, which is designated Shewanella olleyana sp. nov. (type strain ACEM 9(T) = ACAM 644(T) = LMG 21437(T)).

DNA, Bacterial↗