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Light-related plasticity in a variety of crown morphology and within-tree characteristics was examined in sun and shade saplings of Abies amabilis Dougl. ex J. Forbes growing in two late-successional forests with different snow regimes in the Cascade Mountains of Washington, USA. Compared with sun saplings, shade saplings typically had broad flat crowns as a result of acclimation at several scales (needle, shoot, branch, crown and whole sapling). Shoots of shade saplings had a smaller needle mass per unit of stem length than shoots of sun saplings, a feature that enhances light-interception efficiency by reducing among-needle shading. The low annual rate of needle production by shade saplings was associated with a longer needle lifespan and slower needle turnover. Reduced needle production within a shoot was reflected at the branch level, with lateral branches of shade saplings having a smaller needle mass than branches of the same length of sun saplings. Reduced allocation to needles permits greater investment in branches and stems, which is necessary to support the horizontally expanding branch system characteristic of shade saplings. Mean branch age of shade saplings was significantly higher than that of sun saplings. Shade saplings had lower needle mass per unit of trunk biomass or total biomass, reflecting greater investment in the trunk as a support organ. Increased investment in support organs in shade was more evident in the snowier habitat. The observed morphological acclimation makes A. amabilis highly shade and snow-tolerant and thus able to dominate in many late-successional forests in snowy coastal mountain regions.

Nitrogen (N) retranslocation within tree canopies has been intensively studied and assumed to function as a one-way process (e.g., from older to newer leaves). However, recent studies have found that both N output and input occur in individual leaves, suggesting that 'gross' N retranslocation exists behind 'net' N retranslocation. In the present study, the amount and direction of gross N retranslocation within a canopy of deciduous oak Quercus serrata Thunb. ex. Murray saplings were investigated. Labeling was conducted with leaves of Q. serrata saplings cultivated under conditions of low-N (LN) or high-N (HN) fertility. Subsequently, N movement within the canopy was traced. Leaves at two different positions in the canopy (top and lateral) were labeled to determine the direction of gross N retranslocation. To detect seasonal differences, the leaf-labeling experiment was conducted twice during the early and late phases of the growing season. In addition, to compare the quantitative importance of gross N retranslocation and root N uptake, the latter was determined by labeling Q. serrata roots. The N-labeling experiment revealed gross N retranslocation among leaves, i.e., from top to lateral, lateral to top and lateral to lateral positions. Gross N retranslocation was quantitatively more important than root uptake, especially for plants cultivated at LN fertility. Season also affected the amount of gross N retranslocation, and these effects differed between LN and HN fertilities. These findings suggest that N allocation within a canopy is controlled dynamically by both gross N output and input. The mechanisms controlling gross N output and input likely function as key determinants of N allocation within a tree canopy.

Climate change exposes vegetation to unusual levels of drought, risking a decline in productivity and an increase in mortality. It still remains unclear how trees and forests respond to such unusual drought, particularly Southeast Asian tropical rain forests. To understand leaf ecophysiological responses of tropical rain forest trees to soil drying, a rainfall exclusion experiment was conducted on mature canopy trees of Dryobalanops aromatica Gaertn.f. (Dipterocarpaceae) for 4 months in an aseasonal tropical rain forest in Sarawak, Malaysia. The rainfall was intercepted by using a soft vinyl chloride sheet. We compared the three control and three treatment trees with respect to leaf water use at the top of the crown, including stomatal conductance (gsmax), photosynthesis (Amax), leaf water potential (predawn: Ψpre; midday: Ψmid), leaf water potential at turgor loss point (πtlp), osmotic potential at full turgor (π100) and a bulk modulus of elasticity (ε). Measurements were taken using tree-tower and canopy-crane systems. During the experiment, the treatment trees suffered drought stress without evidence of canopy dieback in comparison with the control trees; e.g., Ψpre and Ψmid decreased with soil drying. Minimum values of Ψmid in the treatment trees decreased during the experiment, and were lower than πtlp in the control trees. However, the treatment trees also decreased their πtlp by osmotic adjustment, and the values were lower than the minimum values of their Ψmid. In addition, the treatment trees maintained gs and Amax especially in the morning, though at midday, values decreased to half those of the control trees. Decreasing leaf water potential by osmotic adjustment to maintain gs and Amax under soil drying in treatment trees was considered to represent anisohydric behavior. These results suggest that D. aromatica may have high leaf adaptability to drought by regulating leaf water consumption and maintaining turgor pressure to improve its leaf water relations.

We compared differences in leaf properties, leaf gas exchange and photochemical properties between drought-deciduous and evergreen trees in tropical dry forests, where soil nutrients differed but rainfall was similar. Three canopy trees (Shorea siamensis Miq., Xylia xylocarpa (Roxb.) W. Theob. and Vitex peduncularis Wall. ex Schauer) in a drought-deciduous forest and a canopy tree (Hopea ferrea Lanessan) in an evergreen forest were selected. Soil nutrient availability is lower in the evergreen forest than in the deciduous forest. Compared with the evergreen tree, the deciduous trees had shorter leaf life spans, lower leaf masses per area, higher leaf mass-based nitrogen (N) contents, higher leaf mass-based photosynthetic rates (mass-based P(n)), higher leaf N-based P(n), higher daily maximum stomatal conductance (g(s)) and wider conduits in wood xylem. Mass-based P(n) decreased from the wet to the dry season for all species. Following onset of the dry season, daily maximum g(s) and sensitivity of g(s) to leaf-to-air vapor pressure deficit remained relatively unchanged in the deciduous trees, whereas both properties decreased in the evergreen tree during the dry season. Photochemical capacity and non-photochemical quenching (NPQ) of photosystem II (PSII) also remained relatively unchanged in the deciduous trees even after the onset of the dry season. In contrast, photochemical capacity decreased and NPQ increased in the evergreen tree during the dry season, indicating that the leaves coped with prolonged drought by down-regulating PSII. Thus, the drought-avoidant deciduous species were characterized by high N allocation for leaf carbon assimilation, high water use and photoinhibition avoidance, whereas the drought-tolerant evergreen was characterized by low N allocation for leaf carbon assimilation, conservative water use and photoinhibition tolerance.

Abstract Increases in temperatures and atmospheric CO2 concentration influence the growth performance of trees worldwide. The direction and intensity of tree growth and physiological responses to changing climate do, however, vary according to environmental conditions. Here we present complex, long-term, tree-physiological responses to unprecedented temperature increase in East Asia. For this purpose, we studied radial growth and isotopic (δ13C and δ18O) variations using tree-ring data for the past 100 yr of dominant Quercus mongolica trees from the cool-temperate forests from Hallasan, South Korea. Overall, we found that tree stem basal area increment, intercellular CO2 concentration and intrinsic water-use efficiency significantly increased over the last century. We observed, however, short-term variability in the trends of these variables among four periods identified by change point analysis. In comparison, δ18O did not show significant changes over time, suggesting no major hydrological changes in this precipitation-rich area. The strength and direction of growth–climate relationships also varied during the past 100 yr. Basal area increment (BAI) did not show significant relationships with the climate over the 1924–1949 and 1975–1999 periods. However, over 1950–1974, BAI was negatively affected by both temperature and precipitation, while after 2000, a temperature stimulus was observed. Finally, over the past two decades, the increase in Q. mongolica tree growth accelerated and was associated with high spring–summer temperatures and atmospheric CO2 concentrations and decreasing intrinsic water-use efficiency, δ18O and vapour pressure deficit, suggesting that the photosynthetic rate continued increasing under no water limitations. Our results indicate that the performance of dominant trees of one of the most widely distributed species in East Asia has benefited from recent global changes, mainly over the past two decades. Such findings are essential for projections of forest dynamics and carbon sequestration under climate change.

Fine root respiration is a significant component of carbon cycling in forest ecosystems. Although fine roots differ functionally from coarse roots, these root types have been distinguished based on arbitrary diameter cut-offs (e.g., 2 or 5 mm). Fine root morphology is directly related to physiological function, but few attempts have been made to understand the relationships between morphology and respiration of fine roots. To examine relationships between respiration rates and morphological traits of fine roots (0.15-1.4 mm in diameter) of mature Quercus serrata Murr., we measured respiration of small fine root segments in the field with a portable closed static chamber system. We found a significant power relationship between mean root diameter and respiration rate. Respiration rates of roots<0.4 mm in mean diameter were high and variable, ranging from 3.8 to 11.3 nmol CO2 g(-1) s(-1), compared with those of larger diameter roots (0.4-1.4 mm), which ranged from 1.8 to 3.0 nmol CO2 g(-1) s(-1). Fine root respiration rate was positively correlated with specific root length (SRL) as well as with root nitrogen (N) concentration. For roots<0.4 mm in diameter, SRL had a wider range (11.3-80.4 m g(-1)) and was more strongly correlated with respiration rate than diameter. Our results indicate that a more detailed classification of fine roots<2.0 mm is needed to represent the heterogeneity of root respiration and to evaluate root biomass and root morphological traits.

Allometry of shoot extension units (hereafter termed "current shoots") was analyzed in a Malaysian canopy species, Elateriospermum tapos Bl. (Euphorbiaceae). Changes in current shoot allometry with increasing tree height were related to growth and maintenance of tree crowns. Total biomass, biomass allocation ratio of non-photosynthetic to photosynthetic organs, and wood density of current shoots were unrelated to tree height. However, shoot structure changed with tree height. Compared with short trees, tall trees produced current shoots of the same mass but with thicker and shorter stems. Current shoots with thin and long stems enhanced height growth in short trees, whereas in tall trees, thick and short current shoots may reduce mechanical and hydraulic stresses. Furthermore, compared with short trees, tall trees produced current shoots with more leaves of lower dry mass, smaller area, and smaller specific leaf area (SLA). Short trees adapted to low light flux density by reducing mutual shading with large leaves having a large SLA. In contrast, tall trees reduced mutual shading within a shoot by producing more small leaves in distal than in proximal parts of the shoot stem. The production of a large number of small leaves promoted light penetration into the dense crowns of tall trees. All of these characteristics suggest that the change in current shoot structure with increasing tree height is adaptive in E. tapos, enabling short trees to maximize height growth and tall trees to maximize light capture.

Our knowledge of the nature of belowground competition for moisture and nutrients is limited. In this study, we used an earth impedance method to determine the root absorbing area of Sitka spruce (Picea sitchensis (Bong.) Carr.) trees, making measurements in stands of differing density (2-, 4- and 6-m inter-tree spacing). We compared absorbing root area index (RAI(absorbing); based on the impedance measure) with fine root area index (RAI(fine); based on estimates of total surface area of fine roots) and related these results to investment in conductive roots. Root absorbing area was a near-linear function of tree stem diameter at 1.3 m height. At the stand level, RAI(absorbing), which is analogous to and scaled with transpiring leaf area index (maximum stomatal pore area per unit ground area; LAI(transpiring)), increased proportionally with basal area across the three stands. In contrast, RAI(fine) was inversely propotional to basal area. The ratio of RAI(absorbing) to LAI(transpiring) ranged from 7.7 to 17.1, giving an estimate of the relative aboveground versus belowground resource exchange areas. RAI(absorbing) provides a way of characterizing ecosystem functioning as a physiologically meaningful index of belowground absorbing area.