Responses of Acer saccharum canopy trees and saplings to P, K and lime additions under high N deposition

Heavy atmospheric nitrogen (N) deposition has been associated with altered nutrient cycling, and even N saturation, in forest ecosystems previously thought to be N-limited. This observation has prompted application to such forests of non-N mineral nutrients as a mitigation measure. We examined leaf gas-exchange, leaf chemistry and leaf and shoot morphological responses of Acer saccharum Marsh. saplings and mature trees to experimental additions of non-nitrogenous mineral nutrients (dolomitic lime, phosphorus + potassium (P + K) and lime plus P + K) over 2 years in the Haliburton region of central Ontario, which receives some of the largest annual N inputs in North America. Nutrients were adsorbed in the mineral soil and taken up by A. saccharum trees within 1 year of fertilizer application; however, contrary to expectation, liming had no effect on soil P availability. Saplings and canopy trees showed significant responses to both P + K fertilization and liming, including increased foliar nutrient concentration, leaf size and shoot extension growth; however, no treatment effects on leaf gas-exchange parameters were detected. Increases in shoot extension preceded increases in diameter growth in saplings and canopy trees. Vector analysis of shoot extension growth and nutrient content was consistent with sufficiency of N but marked limitation of P, with co-limitation by calcium (Ca) in saplings and by Ca, Mg and K in canopy trees.     

Water relations and gas exchange of Acer saccharum seedlings in contrasting natural light and water regimes

Field measurements were made of leaf photosynthesis (A), stomatal conductance (g) and leaf water relations for sugar maple (Acer saccharum Marsh.) seedlings growing in a forest understory, small gap or large clearing habitat in southwestern Wisconsin, USA. Predawn water status, leaf gas exchange and plasticity in field and laboratory water relations characteristics were compared among contrasting light environments in a wet year (1987) and a dry year (1988) to evaluate possible interactions between light and water availability in these habitats. Leaf water potentials (Psi(leaf)) at predawn and midday were lower for clearing than gap or understory seedlings. Acclimation of tissue osmotic potentials to light environment was observed among habitats but did not occur within any of the habitats in response to prolonged drought. During a summer drought in 1988, decreases in daily maximum g (g(max)) and maximum A (A(max)) in clearing seedlings were correlated with predawn Psi(leaf), which reached a seasonal minimum of -2.0 MPa. Under well-watered conditions, diurnal fluctuations in Psi(leaf) of up to 2.0 MPa in clearing seedlings occurred along with large midday depressions of A and g. In a wet year, strong stomatal responses to leaf-to-air vapor pressure difference (VPD) in sunny habitats were observed over nine diurnal courses of gas exchange measurements on seedlings in a gap and a clearing. Increasing stomatal limitations to photosynthesis appeared to be responsible for the reduction in A at high VPD for clearing seedlings. In understory seedlings, however, low water-use efficiency and development of leaf water deficits in sunflecks was related to reduced stomatal limitations to photosynthesis relative to seedlings in sunny habitats. Predawn Psi(leaf) and VPD appear to be important factors limiting carbon assimilation in sugar maple seedlings in light-saturating irradiances, primarily through stomatal closure. The overall results are consistent with the idea that sugar maple seedlings exhibit "conservative" water use patterns and have low drought tolerance. Leaf water relations and patterns of water use should be considered in studies of acclimation and species photosynthetic performance in contrasting light environments.     

Seasonal patterns of leaf gas exchange and water relations in dry rain forest trees of contrasting leaf phenology

Diurnal and seasonal patterns of leaf gas exchange and water relations were examined in tree species of contrasting leaf phenology growing in a seasonally dry tropical rain forest in north-eastern Australia. Two drought-deciduous species, Brachychiton australis (Schott and Endl.) A. Terracc. and Cochlospermum gillivraei Benth., and two evergreen species, Alphitonia excelsa (Fenzal) Benth. and Austromyrtus bidwillii (Benth.) Burret. were studied. The deciduous species had higher specific leaf areas and maximum photosynthetic rates per leaf dry mass in the wet season than the evergreens. During the transition from wet season to dry season, total canopy area was reduced by 70-90% in the deciduous species and stomatal conductance (g(s)) and assimilation rate (A) were markedly lower in the remaining leaves. Deciduous species maintained daytime leaf water potentials (Psi(L)) at close to or above wet season values by a combination of stomatal regulation and reduction in leaf area. Thus, the timing of leaf drop in deciduous species was not associated with large negative values of daytime Psi(L) (greater than -1.6 MPa) or predawn Psi(L) (greater than -1.0 MPa). The deciduous species appeared sensitive to small perturbations in soil and leaf water status that signalled the onset of drought. The evergreen species were less sensitive to the onset of drought and g(s) values were not significantly lower during the transitional period. In the dry season, the evergreen species maintained their canopies despite increasing water-stress; however, unlike Eucalyptus species from northern Australian savannas, A and g(s) were significantly lower than wet season values.     

Fruit load and canopy shading affect leaf characteristics and net gas exchange of 'Spring' navel orange trees

Five-year-old 'Spring' navel (Citrus sinensis (L.) Osbeck) orange trees were completely defruited, 50% defruited or left fully laden to study effects of fruit load on concentrations of nitrogen (N) and carbohydrate, net assimilation of CO2 (Ac) and stomatal conductance (gs) of mature leaves on clear winter days just before fruit harvest. Leaves on defruited trees were larger, had higher starch concentrations and greater leaf dry mass per area (LDMa) than leaves on fruited trees. Both Ac and gs were more than 40% lower in sunlit leaves on defruited trees than in sunlit leaves on trees with fruit. Leaves immediately adjacent to fruit were smaller, had lower leaf nitrogen and carbohydrate concentrations, lower LDMa and lower Ac than leaves on non-fruiting branches of the same trees. Removing half the crop increased individual fruit mass, but reduced fruit color development. Half the trees were shaded with 50% shade cloth for 4 months before harvest to determine the effects of lower leaf temperature (Tl) and leaf-to-air vapor pressure difference on leaf responses. On relatively warm days when sunlit Tl > 25 degrees C, shade increased Ac and gs, but had no effect on the ratio of internal to ambient CO2 (Ci/Ca) concentration in leaves, implying that high mesophyll temperatures in sunlit leaves were more important than gs in limiting Ac. Sunlit leaves were more photoinhibited than shaded leaves on cooler days when Tl < 25 degrees C. Shade decreased total soluble sugar concentrations in leaves, but had no effect on leaf starch concentrations. Shading had no effects on canopy volume, yield or fruit size, but shaded fruit developed better external color than sun-exposed fruit. Overall, the presence of a normal fruit crop resulted in lower foliar carbohydrate concentrations and higher Ac compared with defruited trees, except on warm days when Ac was reduced by high leaf temperatures.     

Responses of foliar gas exchange to long-term elevated CO(2) concentrations in mature loblolly pine trees

Branches of field-grown mature loblolly pine (Pinus taeda L.) trees were exposed for 2 years (1992 and 1993) to ambient or elevated CO(2) concentrations (ambient + 165 micro mol mol(-1) or ambient + 330 micro mol mol(-1) CO(2)). Exposure to elevated CO(2) concentrations enhanced rates of net photosynthesis (P(n)) by 53-111% compared to P(n) of foliage exposed to ambient CO(2). At the same CO(2) measurement concentration, the ratio of intercellular to atmospheric CO(2) concentration (C(i)/C(a)) and stomatal conductance to water vapor did not differ among foliage grown in an ambient or enriched CO(2) concentration. Analysis of the relationship between P(n) and C(i) indicated no significant change in carboxylation efficiency of ribulose-1,5-bisphosphate carboxylase/oxygenase during growth in elevated CO(2) concentrations. Based on estimates derived from P(n)/C(i) curves, there were no apparent treatment differences in dark respiration, CO(2) compensation point or P(n) at the mean C(i). In 1992, foliage in the three CO(2) treatments yielded similar estimates of CO(2)-saturated P(n) (P(max)), whereas in 1993, estimates of P(max) were higher for branches grown in elevated CO(2) than in ambient CO(2). We conclude that field-grown loblolly pine trees do not exhibit downward acclimation of leaf-level photosynthesis in their long-term response to elevated CO(2) concentrations.     

Ecophysiological attributes of the native Acer saccharum and the exotic Acer platanoides in urban oak forests in Pennsylvania, USA

When the exotic Acer platanoides L. (Norway maple) and the native A. saccharum Marsh. (sugar maple) grow together in the understories of urban Quercus forests in the eastern USA, average annual height growth increments are nearly twice as large in A. platanoides as in A. saccharum, 19.26 +/- 3.22 versus 10.01 +/- 1.69 cm. We examined several ecophysiological mechanisms that might be associated with the superior invasive ability and growth of A. platanoides in two urban oak forests in Pennsylvania. Leaf longevity was 12 days greater in A. platanoides than in A. saccharum. In addition, leaf mass/leaf area ratio was greater in A. platanoides than in A. saccharum (2.67 +/- 0.03 versus 2.32 +/- 0.02 mg cm(-2)); however, leaf thickness was significantly lower in A. platanoides than in A. saccharum suggesting that A. platanoides contains more dense palisade and mesophyll cell layers than A. saccharum. Field net photosynthesis (mass basis) and photosynthetic light response curves (area basis) indicated significantly greater carbon assimilation, and nitrogen and phosphorus use efficiencies in A. platanoides than in A. saccharum. Acer platanoides also exhibited higher water use efficiency than A. saccharum (0.88 +/- 0.12 versus 0.32 +/- 0.09 mmol CO(2) mol(-1) H(2)O). Acer platanoides exhibited significantly lower osmotic potentials than A. saccharum, but a similar relative water content at zero turgor. We conclude that A. platanoides utilizes light, water and nutrients more efficiently than A. saccharum.     

Effects of regulated deficit irrigation during the pre-harvest period on gas exchange, leaf development and crop yield of mature almond trees

We investigated the effects of regulated deficit irrigation (RDI) during the pre-harvest period (kernel-filling stage) on water relations, leaf development and crop yield in mature almond (Prunus dulcis (Mill.) D.A. Webb cv. Cartagenera) trees during a 2-year field experiment. Trees were either irrigated at full-crop evapotranspiration (ETc=100%) (well-irrigated control treatment) or subjected to an RDI treatment that consisted of full irrigation for the full season, except from early June to early August (kernel-filling stage), when 20% ETc was applied. The severity of water stress was characterized by measurements of soil water content, predawn leaf water potential (Psipd) and relative water content (RWC). Stomatal conductance (gs), net CO2 assimilation rate (A), transpiration rate (E), leaf abscission, leaf expansion rate and crop yield were also measured. In both years, Psipd and RWC of well-irrigated trees were maintained above -1.0 MPa and 92%, respectively, whereas the corresponding values for trees in the RDI treatment were -2.37 MPa and 82%. Long-term water stress led to a progressive decline in gs, A and E, with significant reductions after 21 days in the RDI treatment. At the time of maximum stress (48 days after commencement of RDI), A, gs and E were 64, 67 and 56% lower than control values, respectively. High correlations between A, E and gs were observed. Plant water status recovered within 15 days after the resumption of irrigation and was associated with recovery of soil water content. A relatively rapid and complete recovery of A and gs was also observed, although the recovery was slower than for Psipd and RWC. Severe water stress during the kernel-filling stage resulted in premature defoliation (caused by increased leaf abscission) and a reduction in leaf growth rate, which decreased tree leaf area. Although kernel yield was correlated with leaf water potential, RDI caused a nonsignificant 7% reduction in kernel yield and had no effect on kernel size. The RDI treatment also improved water-use efficiency because about 30% less irrigation water was applied in the RDI treatment than in the control treatment. We conclude that high-cropping almonds can be successfully grown in semiarid regions in an RDI regime provided that Psipd is maintained above a threshold value of -2 MPa.     

Seasonal variation in biomass and carbohydrate partitioning of understory sugar maple (Acer saccharum) and yellow birch (Betula alleghaniensis) seedlings

We investigated seasonal patterns of biomass and carbohydrate partitioning in relation to shoot growth phenology in two age classes of sugar maple (Acer saccharum Marsh.) and yellow birch (Betula alleghaniensis Britt.) seedlings growing in the understory of a partially harvested forest. The high root:shoot biomass ratio and carbohydrate concentration of sugar maple are characteristic of species with truncated growth patterns (i.e., cessation of aboveground shoot growth early in the growing season), a conservative growth strategy and high shade tolerance. The low root:shoot biomass ratio and carbohydrate concentration of yellow birch are characteristic of species with continuous growth patterns, an opportunistic growth strategy and low shade tolerance. In both species, starch represented up to 95% of total nonstructural carbohydrates and was mainly found in the roots. Contrary to our hypothesis, interspecific differences in shoot growth phenology (i.e., continuous versus truncated) did not result in differences in seasonal patterns of carbohydrate partitioning. Our results help explain the niche differentiation between sugar maple and yellow birch in temperate, deciduous understory forests.     

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.     

Determination of fiber length and juvenile and mature wood zones in Acer velutinum Boiss. trees grown in Iran

In our investigation we studied fiber lengths and the transition age from juvenile to mature wood in Acer velutinum Boiss. For this purpose, samples from three normal maple trees at a Noshahr site in northern Iran were selected. Disks were cut at breast height. Test samples were taken along a radial direction from the pith to the bark, accounting for every ring during a 48-year period. We used the Franklin method to distinguish between fibers of juvenile and mature wood. The results show that the fiber length increased along the radial direction from the pith to the bark. The transition age between juvenile and mature wood was determined at the 14th annual ring from the pith.     

Leaf morphology and photosynthetic adjustments among deciduous broad-leaved trees within the vertical canopy profile

Photosynthetic acclimation of deciduous broad-leaved tree species was studied along a vertical gradient within the canopy of a multi-species deciduous forest in northern Japan. We investigated variations in (1) local light regime and CO2 concentration ([CO2]), and (2) morphological (area, thickness and area per mass), biochemical (nitrogen and chlorophyll concentrations) and physiological (light-saturated photosynthetic rate) attributes of leaves of seven major species on three occasions (June, August and October). We studied early successional species, alder (Alnus hirsuta (Spach) Rupr.) and birch (Betula platyphylla var. japonica (Miq.) Hara); gap phase species, walnut (Juglans ailanthifolia Carrière) and ash (Fraxinus mandshurica var. japonica Rupr.); mid-successional species, basswood (Tilia japonica (Miq.) Simonk.) and elm (Ulmus davidiana var. japonica (Rehd.) Nakai); and the late-successional species, maple (Acer mono Bunge). All but maple initiated leaf unfolding from the lower part of the crown. The [CO2] within the vertical profile ranged from 320-350 ppm in the upper canopy to 405-560 ppm near the ground. The lowest and highest ambient [CO2] occurred during the day and during the night, respectively. This trend was observed consistently during the summer, but not when trees were leafless. Chlorophyll concentration was positively related to maximum photosynthetic rate within, but not among, species. Leaf senescence started from the inner part of the crown in alder and birch, but started either in the outer or top portion of the canopy of ash, basswood and maple. Chlorophyll (Chl) to nitrogen ratio in leaves increased with decreasing photon flux density. However, Chl b concentration in all species remained stable until the beginning of leaf senescence. Maximum photosynthetic rates observed in sun leaves of early successional species, gap phase or mid-successional species, and late successional species were 12.5-14.8 micromol m(-2) s(-1), 4.1-7.8 micromol m(-2) s(-1) and 3.1 micromol m(-2) s(-1), respectively.     

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.     

Changes in shoot properties in relation to vertical positions within the crown of mature canopy trees of Abies mariesii and Abies veitchii

We investigated current shoot properties in two contrasting vertical positions (leader crown; LC, and lower branch; LB) within the crowns of mature trees of two subalpine conifer species, Abies mariesii and A. veitchii. For both LCs and LBs, shoot length decreased with increasing branching order. However, shoot properties were different between LCs and LBs. Shoots in LCs had more needle biomass per unit of shoot length. Shoots in sunny conditions pack needles closer along the shoot and intercept incoming light more completely. This causes the shoots in the LCs to have more needles. In contrast, less needle packing per unit shoot length in LBs results in the avoidance of mutual shading among needles in order to intercept limited light more effectively. Because branch systems in lower layers tend to be more shaded, the quantity of irradiance received by the shoots in LBs is smaller. Thus, reduced needle amounts on the shoots in LBs reflect the needle arrangement acclimating to the lower light availability. This study suggests the importance of changes in the properties of individual shoot as a component of a branch system and accordingly a whole-crown system in mature canopy trees of A. mariesii and A. veitchii.     

Influence of canopy position, needle age and season on the foliar gas exchange of Pinus canariensis

We investigated the seasonal variation in the gas exchange of current and 1-year-old needles in the upper sun and lower shade crown of adult Pinus canariensis trees. In general, current year needles displayed lower gas exchange rates than the 1-year-old needles. In both needle age classes, gas exchange was significantly lower in the shade than in the sun crown. However irrespective of crown position and needle age, maximum daily net photosynthesis, transpiration, and stomatal conductance for water vapour were generally higher during the wet and cold winter as compared to the dry and hot summer. These higher gas exchange values obtained during the cold and wet season can mainly be explained by higher soil-water availability and lower evaporative demand as compared to the warm and dry seaon. In addition, we also observed a displacement in the temperature optimum of net photosynthesis towards lower temperatures during the cold and wet season as compared to the warm and dry season. The observed gas exchange characteristics indicate a conservative water saving strategy and thus allowing P. canariensis needles to maintain a positive carbon gain even at periods of high evaporative demand and low soil-water availability.     

Effects of soil and stem base heating on survival, resprouting and gas exchange of Acer and Quercus seedlings

Acer rubrum L., A. saccharum Marsh., Quercus alba L. and Q. rubra L. seedlings subjected to soil and stem base heat treatments showed rapid declines in rates of transpiration and photosynthesis. Reductions in photosynthetic rate were partly attributable to mesophyll inhibition. Quercus seedlings were less able to maintain transpiration and photosynthesis after heat treatment than Acer seedlings. Declines in rates of transpiration and photosynthesis of Quercus seedlings were observed 1 h after heat treatment and became more pronounced over time. In contrast, rates of transpiration and photosynthesis of Acer seedlings initially declined in response to heat treatment, partially recovered after one or two days, but then declined again six to eight days after the heat treatment. Observed changes in leaf water potential after heating were small, suggesting that hydraulic factors were not the primary signal eliciting the gas exchange response to soil and stem heating. Ultimately, the heat treatments caused stem die-back of most seedlings. For all species, seedlings that resprouted had a greater chance of surviving heat stress than seedlings that did not resprout. Despite the rapid loss of photosynthetic capacity in response to heat treatment in Quercus seedlings, survival was higher in Quercus seedlings than in Acer seedlings, and was associated with a greater capacity for resprouting. We suggest that the reduced allocation of resources toward recovery of photosynthesis in existing Quercus stems after heat stress is a physiological mechanism that facilitates resprouting and hence survival of Quercus seedlings after fire.     

Seasonal patterns of radial root growth and starch dynamics in plantation-grown Sitka spruce trees of different ages

Seasonal patterns of radial root growth within 1 m of tree stems were examined in Scottish plantations of Sitka spruce trees aged 9, 15 and 20 years. Results were compared with parallel measurements of shoot extension, radial growth of stems and amounts of starch stored in tissues external to root wood. Youngest trees produced the largest annual increments in root cross-sectional area and numbers of new cells along radial files of tracheids. Irrespective of tree age, new cells were present in roots before bud burst and the onset of radial growth occurred progressively later with increasing distances from the stems. At ages 15 and 20, both stem cross-sectional area and radial root growth up to 0.5 m from the stem base had a minor peak of activity preceding and a major peak following shoot elongation. Further than 0.5 m from the stem, root growth was frequently restricted to the period following shoot extension. Starch storage in the roots reached a maximum in April and May, which was greatest for 9-year-old trees and least for 20-year-old trees. At all ages, radial root growth in early spring occurred concurrently with increased starch storage. Later in the season starch reserves declined rapidly during the period of shoot elongation and root growth occurred whilst reserves were low. At all ages for positions on the root at the base of the stem and 0.25 m from it, starch depletion, at its maximum rate during June, accounted for less than the measured increment of root wood growth at that point. This indicates a substantial translocation of substrates to these zones during growth. At the same time, the reduction in starch concentrations at more distal points from the stem far exceeded that required for local root thickening.     

Seasonal photosynthetic gas exchange and leaf reflectance characteristics of male and female cottonwoods in a riparian woodland

Cottonwoods (Populus spp.) are dioecious phreatophytes of hydrological and ecological importance in riparian woodlands throughout the Northern Hemisphere. In streamside zones of southern Alberta, groundwater and soil water typically decline between May and September. To understand how narrowleaf cottonwoods (Populus angustifolia James) are adapted to this seasonal decrease in water availability, we measured photosynthetic gas exchange, leaf reflectance, chlorophyll fluorescence and stable carbon isotope composition (delta(13)C) in trees growing in the Oldman River valley of southern Alberta during the 2006 growth season. Accompanying the seasonal recession in river flow, groundwater table depth (Z(gw)) declined by 1.6 m, but neither mean daily light-saturated net photosynthetic rate (A(max)) nor stomatal conductance (g(s)) was correlated with this change. Both A(max) and g(s) followed a parabolic seasonal pattern, with July 24 maxima of 15.8 micromol m(-2) s(-1) and 559 mmol m(-2) s(-1), respectively. The early summer rise in A(max) was related to an increase in the chlorophyll pool during leaf development. Peak A(max) coincided with the maximum quantum efficiency of Photosystem II (F(v)/F(m)), chlorophyll index (CI) and scaled photochemical reflectance index (sPRI), but occurred one month after maximum volumetric soil water (theta(v)) and minimum Z(gw). In late summer, A(max) decreased by 30-40% from maximum values, in weak correlation with theta(v) (r(2) = 0.50). Groundwater availability limited late-season water stress, so that there was little variation in mean daily transpiration (E). Decreasing leaf nitrogen (% dry mass), CI, F(v)/F(m) and normalized difference vegetation index (NDVI) were also consistent with leaf aging effects. There was a strong correlation between A(max) and g(s) (r(2) = 0.89), so that photosynthetic water-use efficiency (WUE; A(max)/E) decreased logarithmically with increasing vapor pressure deficit in both males (r(2) = 0.75) and females (r(2) = 0.95). The male:female ratio was unequal (2:1, chi(2) = 16.5, P < 0.001) at the study site, but we found no significant between-sex differences in photosynthetic gas exchange, leaf reflectance or chlorophyll fluorescence that might explain the unequal ratio. Females tended to display lower NDVI than males (P = 0.07), but mean WUE did not differ significantly between males and females (2.1 +/- 0.2 versus 2.5 +/- 0.2 mmol mol(-1)), and delta(13)C remained in the -28.8 to -29.3 per thousand range throughout the growth season, in both sexes. These results demonstrate changes in photosynthetic and water-use characteristics that collectively enable vigorous growth throughout the season, despite seasonal changes in water supply and demand.     

Growth form and seasonal variation in leaf gas exchange of Colophospermum mopane savanna trees in northwest Botswana

We investigated differences in physiological and morphological traits between the tall and short forms of mopane (Colophospermum mopane (Kirk ex Benth.) Kirk ex J. Léonard) trees growing near Maun, Botswana on a Kalahari sandveld overlying an impermeable calcrete duricrust. We sought to determine if differences between the two physiognomic types are attributable to the way they exploit available soil water. The tall form, which was located on deeper soil than the short form (5.5 versus 1.6 m), had a lower leaf:fine root biomass ratio (1:20 versus 1:6), but a similar leaf area index (0.9-1.0). Leaf nitrogen concentrations varied between 18 and 27 mg g(-1) and were about 20% higher in the tall form than in the short form. Maximum net assimilation rates (A sat) occurred during the rainy seasons (March-April 2000 and January-February 2001) and were similar in the tall and short forms (15-22 micromol m(-2) s(-1)) before declining to less than 10 micromol m(-2) s(-1) at the end of the rainy season in late April. As the dry season progressed, A sat, soil water content, predawn leaf water potential (Psi pd) and leaf nitrogen concentration declined rapidly. Before leaf abscission, Psi pd was more negative in the short form (-3.4 MPa) than in the tall form (-2.7 MPa) despite the greater availability of soil water beneath the short form trees. This difference appeared attributable to differences in root depth and density between the physiognomic types. Stomatal regulation of water use and carbon assimilation differed between years, with the tall form having a consistently more conservative water-use strategy as the dry season progressed than the short form.     

枫香人工林枯落物和土壤层重金属含量的季节变化

对长沙市郊枫香人工林枯落物和土壤层重金属含量季节变化进行研究。结果表明:枯落物中Fe、Zn、Ni含量季节之间的变化差异性显著(P0.05),Mn、Cu、Pb、Cd元素含量季节之间的变化差异性不显著(P0.05);7种重金属元素含量在A、B、C层土壤之间差异性和每层季节变化之间的差异性都不显著(P0.05);枯落物与A层土壤重金属含量的季节变化正好相反,A层土壤重金属含量春季到秋季逐步增加,秋季达到最大值,冬季回落。     

Diurnal changes in leaf gas exchange characteristics in the uppermost canopy of a rain forest tree, Dryobalanops aromatica Gaertn. f

Dryobalanops aromatica Gaertn. f. is a major tropical canopy species in lowland tropical rain forests in Peninsular Malaysia. Diurnal changes in net photosynthetic rate (A) and stomatal conductance to water vapor (g(s)) were measured in fully expanded young and old leaves in the uppermost canopy (35 m above ground). Maximum A was 12 and 10 micro mol m(-2) s(-1) in young and old leaves, respectively; however, because of large variation in A among leaves, mean maximum A in young and old leaves was only 6.6 and 5.5 micro mol m(-2) s(-1), respectively. Both g(s) and A declined in young leaves when T(leaf) exceeded 34 degrees C and leaf-to-air vapor pressure deficit (DeltaW) exceeded 0.025, whereas in old leaves, g(s) and A did not start to decline until T(leaf) and DeltaW exceeded 36 degrees C and 0.035, respectively. Under saturating light conditions, A was linearly related to g(s). The coefficient of variation (CV) for the difference between the CO(2) concentrations of ambient air and the leaf intercellular air space (C(a) - C(i)) was smaller than the CV for A or g(s), suggesting that maximum g(s) was mainly controlled by mesophyll assimilation (A/C(i)). Minimum C(i)/C(a) ratios were relatively high (0.72-0.73), indicating a small drought-induced stomatal limitation to A and non-conservative water use in the uppermost canopy leaves.