Leaf conductance and CO(2) assimilation of Larix gmelinii growing in an eastern Siberian boreal forest

In July 1993, we measured leaf conductance, carbon dioxide (CO(2)) assimilation, and transpiration in a Larix gmelinii (Rupr.) Rupr. ex Kuzen forest in eastern Siberia. At the CO(2) concentration of ambient air, maximum values (mean of 10 highest measured values) for CO(2) assimilation, transpiration and leaf conductance for water vapor were 10.1 micro mol m(-2) s(-1), 3.9 mmol m(-2) s(-1) and 365 mmol m(-2) s(-1), respectively. The corresponding mean values, which were much lower than the maximum values, were 2.7 micro mol m(-2) s(-1), 1.0 mmol m(-2) s(-1) and 56 mmol m(-2) s(-1). The mean values were similar to those of Vaccinium species in the herb layer. The large differences between maximum and actual performance were the result of structural and physiological variations within the tree crowns and between trees that reduced maximum assimilation and leaf conductance by about 40 and 60%, respectively. Thus, maximum assimilation and conductance values averaged over the canopy were 6.1 micro mol m(-2) s(-1) and 146 mmol m(-2) s(-1), respectively. Dry air caused stomatal closure, which reduced assimilation by an additional 26%. Low irradiances in the morning and evening had a minor effect (-6%). Daily canopy transpiration was estimated to be 1.45 mm day(-1), which is higher than the value of 0.94 mm day(-1) measured by eddy covariance, but similar to the value of 1.45 mm day(-1) calculated from the energy balance and soil evaporation, and less than the value of 2.1 mm day(-1) measured by xylem flux. Daytime canopy carbon assimilation, expressed on a ground area basis, was 0.217 mol m(-2) day(-1), which is higher than the value measured by eddy flux (0.162 mol m(-2) day(-1) including soil respiration). We discuss the regulation of leaf gas exchange in Larix under the extreme climatic conditions of eastern Siberia (temperature > 35 degrees C and vapor pressure deficit > 5.0 kPa).     

Root turnover and relocation in the soil profile in response to seasonal soil water variation in a natural stand of Utah juniper (Juniperus osteosperma)

Juniper species are noted for long-lived foliage, low and persistent gas exchange activity and drought tolerance. Because leaves and roots of the same species are thought to be similar in structure and life history, we hypothesized that Juniperus osteosperma (Torr.) Little (Utah juniper) fine roots would reflect the persistent aboveground foliage characteristic of this species. We monitored fine roots, less than 1 mm in diameter, by minirhizotron imaging to a depth of 150 cm over two growing seasons from April 2002 to December 2003. We measured fine root numbers, lengths and diameters, and noted the time of birth and death of root segments. We correlated our root data with soil water potential measured by thermocouple psychrometry and ecosystem evapotranspiration measured by ecosystem eddy flux. Median fine root lifespan, determined by the Kaplan-Meier product-limit method, was about one year, much less than foliage lifespan estimates of more than five years. Yet, roots of juniper live much longer than those of other Great Basin species. The median survivorship of shallow and deep roots was 144 and 448 days, respectively. Production of new roots was observed during periods of favorable soil water potential and there was a seasonal progression of increased new roots and root length during the warm season toward lower soil depths with root loss in the upper soil layers. This was also reflected in water extraction which progressed to greater soil depths later in the warm season. Aboveground, rates of ecosystem evapotranspiration decreased with decreasing soil water potentials in a similar manner in both 2002 and 2003, reflecting the relocation of roots to available water at depth. Juniper exhibited a flexible root depth distribution throughout the 20 months of this study, indicating the potential to respond to shifting soil water resources despite long fine root lifespans.     

Gas exchange and carbon isotope discrimination of Juniperus osteosperma and Juniperus occidentalis across environmental gradients in the Great Basin of western North America

We determined how ecophysiological characteristics of two juniper species, Juniperus occidentalis Hook. (western juniper) and Juniperus osteosperma (Torr.) Little (Utah juniper), changed along altitudinal and regional environmental gradients in the Great Basin of western North America. We obtained diurnal measurements of leaf gas exchange and xylem water potential (Psi) from plants at a low and a high altitude site within each of six mountain ranges during fall 1994, spring, summer, and fall 1995, and summer 1996. We also determined carbon isotope composition (delta(13)C) of leaf cellulose produced during the 1995 growing season. Overall, leaf gas exchange, Psi and delta(13)C did not differ significantly between species. Differences in daily (A(d)) and season-long (A(s)) carbon assimilation among mountain ranges suggested two groupings-a group of northern ranges and a group of southern ranges. Each group contained one mountain range with J. occidentalis and two with J. osteosperma. Differences in carbon assimilation based on this grouping were associated with two findings: (1) conductance of CO(2) from substomatal cavities to the site of carboxylation (g(m)) for junipers in the northern ranges averaged almost twice that of junipers in the southern ranges; and (2) physiological shifts occurred such that A(d) of junipers in the northern ranges was influenced more by Psi(pd), whereas A(d) of junipers in the southern ranges was influenced more by leaf temperature. Mean delta(13)C over all trees at a site was significantly correlated with annual precipitation. Significant differences in A(d) occurred between altitudes, but these differences were associated with differences in the timing of optimum leaf temperature for photosynthesis rather than with physiological acclimation to temperature, irradiance, or Psi. Most gas exchange parameters (e.g., assimilation, transpiration, stomatal conductance, and water use efficiency) varied seasonally, and the seasonal differences were strongly influenced by water stress.     

Transpiration by two poplar varieties grown as coppice for biomass production

Fast-growing tree clones selected for biomass plantations are highly productive and therefore likely to use more water than the agricultural crops they replace. We report field measurements of transpiration through the summer of 1994 from two poplar clones, Beaupré (Populus trichocarpa Torr. & A. Gray x P. deltoides Bartr. ex Marsh.) and Dorschkamp (P. deltoides x P. nigra L.), grown as unirrigated short-rotation coppice in southern England. Stand transpiration was quantified by scaling up from sap flow measurements made with the heat balance method in a sample of stems. Leaf conductances, leaf area development, meteorological variables and soil water deficit were also measured to investigate the response of the trees to the environment. High rates of transpiration were found for Beaupré. In June, when soil water was plentiful, the mean (+/- SD) transpiration rate over an 18-day period was 5.0 +/- 1.8 mm day(-1), reaching a maximum of 7.9 mm day(-1). Transpiration rates from Dorschkamp were lower, as a result of its lower leaf area index. High total leaf conductances were measured for both Beaupré (0.34 +/- 0.17 mol m(-2) s(-1)) and Dorschkamp (0.39 +/- 0.16 mol m(-2) s(-1)). Leaf conductance declined slightly with increasing atmospheric vapor pressure deficit in both clones, but only in Beaupré did leaf conductance decrease as soil water deficit increased.     

Water stress decreases the transfer conductance of Douglas-fir (Pseudotsuga menziesii) seedlings

We tested the hypothesis that transfer conductance (gi) of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings is reduced by water stress. Seedlings were irrigated with a solution of 25% polyethylene glycol so as to impose water stress rapidly, thereby limiting acclimatory responses. Transfer conductance was measured pre-treatment and post-treatment by two methods. Water stress reduced net photosynthesis by 20-50%. The initial slope of the rate of photosynthesis (A) over the intercellular carbon dioxide (CO2) concentration (Ci) response was reduced by water stress, indicating that reduced photosynthesis was not wholly accounted for by reduced stomatal conductance. The carbon isotope and chlorophyll fluorescence methods both indicated that water stress decreased gi. From isotopic measurements with 1% O2, gi was 0.076 +/- 0.009 (mean +/- SE) mol m(-2) s(-1) in well-watered seedlings and 0.044 +/- 0.004 mol m(-2) s(-1) in water-stressed seedlings. Fluorescence estimates of gi were 0.08 +/- 0.01 mol m(-2) s(-1) in well-watered seedlings and 0.044 +/- 0.004 mol m(-2) s(-1) in water-stressed seedlings. The drought-induced reduction in gi was responsible for the reduction in slope of the A/Ci response, and thus there was no difference in the slope of the A over the chloroplastic CO2 concentration (Cc) response between treatments and no indication of impaired mesophyll metabolism. These data illustrate that impairments of mesophyll metabolism can be revealed only from analysis of the A/Cc response.     

Seasonal and topographic patterns of forest floor CO(2) efflux from an upland oak forest

Forest floor CO(2) efflux (FF(cer)) is an important component of global carbon budgets, but the spatial variability of forest floor respiration within a forest type is not well documented. Measurements of FF(cer) were initiated in mid-March of 1991 and continued at biweekly to monthly intervals until mid-November. Observations were made at 45 sites along topographic gradients of the Walker Branch Watershed, Tennessee including northeast and southwest facing slopes, valley-bottoms, and exposed ridge-top locations. The FF(cer) measurements were made with a portable gas-exchange system, and all observations were accompanied by soil temperature and soil water content measurements. As expected, FF(cer) exhibited a distinct seasonal trend following patterns of soil temperature, but soil water content and the volume percent of the soil's coarse fraction were also correlated with observed rates. Over the entire measurement period, FF(cer) ranged from a typical minimum of 0.8 micro mol m(-2) s(-1) to an average maximum near 5.7 micro mol m(-2) s(-1). No significant differences in FF(cer) were observed among the ridge-top and slope positions, but FF(cer) in the valley-bottom locations was lower on several occasions. An empirical model of FF(cer) based on these observations is suggested for application to whole-stand estimates of forest carbon sequestration.     

Tree water use and rainfall partitioning in a mature poplar-pasture system

Traditionally, poplars (Populus) have been planted to control erosion on New Zealand's hill-slopes, because of their capacity to dry out and bind together the soil, by reducing effective rainfall and increasing evapotranspiration and soil strength. However, the effect of widely spaced poplars on the partitioning of soil water and rainfall has not been reported. This study determined rainfall partitioning for 18 mid-spring days in a mature P. deltoides (Bart. ex Marsh, Clone I78)-pasture association (37 stems per hectare, unevenly spaced at 16.4 +/- 0.4 m) and compared it with a traditional open pasture system in grazed areas of a hill environment. Tree transpiration was measured by the heat pulse technique. A time-driven mathematical model was used to set a zero offset, adjust anomalous values and describe simultaneous sap velocity time courses of trees. The model showed that daylight sap flow velocities can be represented with a nonlinear Beta function (R(2) > 0.98), and differences in the parameters representing the initiation, duration and conformation of the sap velocity can be tested statistically to discern tree transpiration differences during the day. Evapotranspiration was greater for the poplar-pasture association than for the open pasture (2.7-3.0 versus 2.2 mm day(-1)). The tree canopy alone contributed 0.92 mm day(-1) as transpiration and 1.37 mm day(-1) as interception, whereas evapotranspiration of the pasture understory was only 0.4-0.6 mm day(-1). Despite the higher water use of the poplar-pasture association, soil water in the 0-300 mm soil stratum was higher than, or similar to, that of the open pasture. Tree shading decreased evapotranspiration and pasture accumulation under the trees.     

Leaf physiology, production, water use, and nitrogen dynamics of the grassland invader Acacia smallii at elevated CO(2) concentrations

Invasion by woody legumes can alter hydrology, nutrient accumulation and cycling, and carbon sequestration on grasslands. The rate and magnitude of these changes are likely to be sensitive to the effects of atmospheric CO(2) enrichment on growth and water and nitrogen dynamics of leguminous shrubs. To assess potential effects of increased atmospheric CO(2) concentrations on plant growth and acquisition and utilization of water and nitrogen, seedlings of Acacia smallii Isely (huisache) were grown for 13 months at CO(2) concentrations of 385 (ambient), 690, and 980 micro mol mol(-1). Seedlings grown at elevated CO(2) concentrations exhibited parallel declines in leaf N concentration and photosynthetic capacity; however, at the highest CO(2) concentration, biomass production increased more than 2.5-fold as a result of increased leaf photosynthetic rates, leaf area, and N(2) fixation. Measurements of leaf gas exchange and aboveground biomass production and soil water balance indicated that water use efficiency increased in proportion to the increase in atmospheric CO(2) concentration. The effects on transpiration of an accompanying decline in leaf conductance were offset by an increase in leaf area, and total water loss was similar across CO(2) treatments. Plants grown at elevated CO(2) fixed three to four times as much N as plants grown at ambient CO(2) concentration. The increase in N(2) fixation resulted from an increase in fixation per unit of nodule mass in the 690 micro mol mol(-1) CO(2) treatment and from a large increase in the number and mass of nodules in plants in the 980 micro mol mol(-1) CO(2) treatment. Increased symbiotic N(2) fixation by woody invaders in response to CO(2) enrichment may result in increased N deposition in litterfall, and thus increased productivity on many grasslands.     

Land-use changes alter CO2 flux patterns of a tall-grass Andropogon field and a savanna-woodland continuum in the Orinoco lowlands

Land use changes in the savannas of the Orinoco lowlands have resulted in a mosaic of vegetation. To elucidate how these changes have affected carbon exchanges with the atmosphere, we measured CO2 fluxes by eddy covariance and soil CO2 efflux systems along a disturbance gradient beginning with a cultivated tall-grass Andropogon field (S1) and extending over three savanna sites with increasing woody cover growing above native herbaceous vegetation. The savanna sites included a herbaceous savanna (S2), a tree savanna (S3) and a woodland savanna (S4). During the wet season, maximum diurnal net ecosystem exchange (NEE) over the S1-S4 sites was 6.6-9.3, 6.6-7.9, 10.6-11.3 and 9.3-10.6 micromol m(-2) s(-1), respectively. The rate of CO2 uptake over S1 was lower than that for C4 grasses elsewhere because of pasture degradation. Soil respiration and temperature were exponentially related when soil water content (theta) was above 0.083 m(3) m(-3); however, soil respiration declined markedly as theta decreased from 0.083-0.090 to 0.033-0.056 m(3) m(-3). There were bursts of CO2 emission when dry soils were rewetted by rainfall. During the wet season, all sites constituted carbon sinks with maximum net daily ecosystem production (NEP) of 2.1, 1.7, 2.1 and 2.1 g C m(-2) day(-1), respectively. During the dry season, the savanna sites (S2-S4) became carbon sources with maximum emission fluxes of -0.5, -1.4 and -1.6 g C m(-2) day(-1), respectively, whereas the tall-grass field (S1) remained a carbon sink with a maximum NEP of 0.3 g C m(-2) day(-1) at the end of the season. For all measurement periods, annual NEP of sites S1-S4 was 366, 6, 116 and 139 g C m(-2), respectively. Comparisons of carbon source/sink dynamics across a wide range of savannas indicate that savanna carbon budgets can change in sign and magnitude. On an annual basis, gross primary production over the S1-S4 stands was 797, 803, 136 and 1230 g C m(-2), respectively. Net primary productivity (NPP) of the S1-S4 stands, calculated from eddy covariance measurements as the daily sum of NEE and day and night heterotrophic respiration was 498, 169, 181 and 402 g C m-2 year-1, respectively. These values were slightly higher than NPP based on harvest measurements (432, 162, 176 and 386 g C m(-2) year(-1), respectively), presumably because fine roots were incompletely harvested. Soil water content limited carbon uptake at all sites, and water-use efficiency (WUE) was related to rainfall dynamics. During the dry season, all sites except the cultivated tall-grass Andropogon field (S1) had a negative WUE. Although our results are specific to the Orinoco vegetational mosaic, the effects of land-use practices on the controls and physiological functions of the studied ecosystems may be generalized to other savannas.     

Characteristics of photosynthesis and stomatal conductance in the shrubland species manuka (Leptospermum scoparium) and kanuka (Kunzea ericoides) for the estimation of annual canopy carbon uptake

Responses of photosynthesis to carbon dioxide (CO2) partial pressure and irradiance were measured on leaves of 39-year-old trees of manuka (Leptospermum scoparium J. R. Forst. & G. Forst.) and kanuka (Kunzea ericoides var. ericoides (A. Rich.) J. Thompson) at a field site, and on leaves of young trees grown at three nitrogen supply rates in a nursery, to determine values for parameters in a model to estimate annual net carbon uptake. These secondary successional species belong to the same family and commonly co-occur. Mean (+/- standard error) values of the maximum rate of carboxylation (hemi-surface area basis) (Vcmax) and the maximum rate of electron transport (Jmax) at the field site were 47.3 +/- 1.9 micromol m(-2) s(-1) and 94.2 +/- 3.7 micromol m(-2) s(-1), respectively, with no significant differences between species. Both Vcmax and Jmax were positively related to leaf nitrogen concentration on a unit leaf area basis, and the slopes of these relationships did not differ significantly between species or between the trees in the field and young trees grown in the nursery. Mean values of Jmax/Vcmax measured at 20 degrees C were significantly lower (P < 0.01) for trees in the field (2.00 +/- 0.05) than for young trees in the nursery with similar leaf nitrogen concentrations (2.32 +/- 0.08). Stomatal conductance decreased sharply with increasing air saturation deficit, but the sensitivity of the response did not differ between species. These data were used to derive parameters for a coupled photosynthesis-stomatal conductance model to scale estimates of photosynthesis from leaves to the canopy, incorporating leaf respiration at night, site energy and water balances, to estimate net canopy carbon uptake. Over the course of a year, 76% of incident irradiance (400-700 nm) was absorbed by the canopy, annual net photosynthesis per unit ground area was 164.5 mol m(-2) (equivalent to 1.97 kg C m(-2)) and respiration loss from leaves at night was 37.5 mol m(-2) (equivalent to 0.45 kg m(-2)), or 23% of net carbon uptake. When modeled annual net carbon uptake for the trees was combined with annual respiration from the soil surface, estimated net primary productivity for the ecosystem (0.30 kg C m(-2)) was reasonably close to the annual estimate obtained from independent mensurational and biomass measurements made at the site (0.22 +/- 0.03 kg C m(-2)). The mean annual value for light-use efficiency calculated from the ratio of net carbon uptake (net photosynthesis minus respiration of leaves at night) and absorbed irradiance was 13.0 mmol C mol(-1) (equivalent to 0.72 kg C GJ(-1)). This is low compared with values reported for other temperate forests, but is consistent with limitations to photosynthesis in the canopy attributable mainly to low nitrogen availability and associated low leaf area index.     

Whole-plant water flux in understory red maple exposed to altered precipitation regimes

Sap flow gauges were used to estimate whole-plant water flux for five stem-diameter classes of red maple (Acer rubrum L.) growing in the understory of an upland oak forest and exposed to one of three large-scale (0.64 ha) manipulations of soil water content. This Throughfall Displacement Experiment (TDE) used subcanopy troughs to intercept roughly 30% of the throughfall on a "dry" plot and a series of pipes to move this collected precipitation across an "ambient" plot and onto a "wet" plot. Saplings with a stem diameter larger than 10 cm lost water at rates 50-fold greater than saplings with a stem diameter of 1 to 2 cm (326 versus 6.4 mol H(2)O tree(-1) day(-1)). These size-class differences were driven largely by differences in leaf area and cross-sectional sapwood area, because rates of water flux expressed per unit leaf area (6.90 mol H(2)O m(-2) day(-1)) or sapwood area (288 mol H(2)O dm(-2) day(-1)) were similar among saplings of the five size classes. Daily and hourly rates of transpiration expressed per unit leaf area varied throughout much of the season, as did soil matrix potentials, and treatment differences due to the TDE were observed during two of the seven sampling periods. On July 6, midday rates of transpiration averaged 1.88 mol H(2)O m(-2) h(-1) for saplings in the "wet" plot, 1.22 mol H(2)O m(-2) h(-1) for saplings in the "ambient" plot, and 0.76 mol H(2)O m(-2) h(-1) for saplings in the "dry" plot. During the early afternoon of August 28, transpiration rates were sevenfold lower for saplings in the "dry" plot compared to saplings in the "wet" plot and 2.5-fold lower compared to saplings in the "ambient" plot. Treatment differences in crown conductance followed a pattern similar to that of transpiration, with values that averaged 60% lower for saplings in the "dry" plot compared to saplings in the "wet" plot and 35% lower compared to saplings in the "ambient" plot. Stomatal and boundary layer conductances were roughly equal in magnitude. Estimates of the decoupling coefficient (Omega) ranged between 0.64 and 0.72 for saplings in the three TDE treatment plots. We conclude that red maple saplings growing in the understory of an upland oak forest are responsive to their edaphic and climatic surroundings, and because of either their small stature or their shallow root distribution, or both, are likely to be impacted by precipitation changes similar to those predicted by global climate models.     

Effects of short-term ozone exposure and soil water availability on the carbon economy of juvenile Douglas-fir

Effects of ozone and soil water availability on partitioning and translocation of assimilates were studied in three-year-old Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) seedlings exposed, in separate experiments, to 0 and 106 or 0 and 514 micro g m(-3) ozone for 8 h day(-1) for 9 days. The dynamics of carbon from assimilated (14)CO(2) were followed. No interactions between ozone and soil water content were observed. Total net uptake of carbon was reduced by low soil water content, but was unaffected by ozone. Both ozone and low soil water content increased the amount of (14)C-photosynthates retained in the current-year needles. Total starch content in old and current-year needles was unaffected by ozone, but was reduced by low water availability. Translocation of carbon to the root-soil compartment was additively affected by ozone and low soil water content. The results suggest that dry periods in summer combined with high ozone concentrations cause the greatest reduction in the supply of carbon compounds to the root-soil compartment.     

Water deficits at anthesis reduce CO(2) assimilation and yield of lychee (Litchi chinensis Sonn.) trees

Ten-year-old 'Tai So' lychee (Litchi chinensis Sonn.) trees growing on a sandy loam soil in subtropical South Africa (latitude 25 degrees S) were watered weekly (well-watered treatment) or droughted from late July until January (drought treatment). After 16 weeks, at which time the trees obtained most of their water from below 150 cm, average soil water content at 0 to 150 cm depth was 14.5 +/- 0.1% in the well-watered treatment and reached a minimum of 7.6% in the drought treatment. At Week 7, minimum leaf water potential (Psi(L)) in the morning and early afternoon declined to -2.6 and -2.8 MPa, respectively, in droughted trees compared with -1.5 and -2.2 MPa, respectively, in well-watered trees. From Week 9, stomatal conductance and net CO(2) assimilation rate ranged from 70 to 300 mmol m(-2) s(-1) and 3 to 13 micro mol CO(2) m(-2) s(-1), respectively, in well-watered trees. The corresponding values for droughted trees were 50 to 180 mmol m(-2) s(-1) and 2 to 6 micro mol CO(2) m(-2) s(-1). Five weeks after rewatering the droughted trees, gas exchange had not recovered to the rate in well-watered trees, although tree water status recovered within a week of rewatering. In the well-watered trees, water use (E(t)) was 26 +/- 1 mm week(-1) with evaporation (E(p)) of 20 to 70 mm week(-1) indicating a crop factor (k(c) = E(t)/E(p)) of 0.4 to 1.2. Before anthesis, tree water status did not affect extension growth of floral panicles or leafy shoots. In contrast, no vegetative shoots were initiated after fruit set in the droughted trees when Psi(L) in the morning declined to -2.5 MPa. Water deficits reduced initial fruit set by 30% and final fruit set by 70% as a result of fruit splitting (41.2 +/- 4.0% versus 10.0 +/- 1.3%). Water deficits did not alter the sigmoidal pattern of fruit growth, but reduced yield from 51.4 +/- 5.5 kg tree(-1) in well-watered trees to 7.4 +/- 3.3 kg tree(-1) in droughted trees.     

Quantification of two-year-old hybrid poplar root systems: morphology, biomass, and (14)C distribution

Root morphology, biomass, and (14)C distribution were studied in two 2-year-old Populus trichocarpa x P. deltoides hybrids, which originated from hardwood cuttings, to determine the pattern of root distribution in a plantation and to refine methods for root recovery. The trees were labeled with (14)CO(2) and harvested after a 72-hour chase period. Roots attached to each labeled tree were analyzed for morphological traits at the time of harvest. Detached roots from within a 1-m(3) volume of soil surrounding each tree were separated from the soil and sorted on the basis of rooting depth and root diameter. Lateral roots > 2 mm in diameter had a largely horizontal orientation at their point of origin from the cutting and extended horizontally up to 4 m from the cutting. This resulted in considerable overlap of root systems in the plantation. Results from (14)C labeling indicated that 24 +/- 4% (+/- SD) of the carbon exported from branches-labeled within two weeks after branch budset-was translocated to the root system. Dilution of the root (14)C label indicated that from 0 (> 5 mm diameter roots) to 75% (< 2 mm diameter roots) of the roots recovered from within the 1-m(3) volume of soil surrounding a harvested tree originated from other trees. Total root biomass was 6 +/- 1 Mg ha(-1) for both hybrids. Sixty percent of the root biomass was recovered directly from excavation, 16% from coarse-sieving excavated soil, and 24% from re-sorting sieved soil. The study indicated that root growth of hybrid poplars may be rapid and extensive and that detailed sorting of soil subsamples substantially improves the recovery of fine roots < 2 mm in diameter.     

Leaf gas exchange of understory spruce-fir saplings in relict cloud forests, southern Appalachian Mountains, USA

The southern Appalachian spruce-fir (Picea rubens Sarg. and Abies fraseri (Pursh) Poir.) forest is found only on high altitude mountain tops that receive copious precipitation ( > 2000 mm year(-1)) and experience frequent cloud immersion. These high-elevation, temperate rain forests are immersed in clouds on approximately 65% of the total growth season days and for 30-40% of a typical summer day, and cloud deposition accounts for up to 50% of their annual water budget. We investigated environmental influences on understory leaf gas exchange and water relations at two sites: Mt. Mitchell, NC (MM; 35 degrees 45'53' N, 82 degrees 15'53' W, 2028 m elevation) and Whitetop Mtn., VA (WT; 36 degrees 38'19' N, 81 degrees 36'19' W, 1685 m elevation). We hypothesized that the cool, moist and cloudy conditions at these sites exert a strong influence on leaf gas exchange. Maximum photosynthesis (A(max)) varied between 1.6 and 4.0 micromol CO(2) m(-2) s(-1) for both spruce and fir and saturated at irradiances between approximately 200 and 400 micromol m(-2) s(-1) at both sites. Leaf conductance (g) ranged between 0.05 and 0.25 mol m(-2) s(-1) at MM and between 0.15 and 0.40 mol m(-2) s(-1) at WT and was strongly associated with leaf-to-air vapor pressure difference (LAVD). At both sites, g decreased exponentially as LAVD increased, with an 80-90% reduction in g between 0 and 0.5 kPa. Predawn leaf water potentials remained between -0.25 and -0.5 MPa for the entire summer, whereas late afternoon values declined to between -1.25 and -1.75 MPa by late summer. Thus, leaf gas exchange appeared tightly coupled to the response of g to LAVD, which maintained high water status, even at the relatively low LAVD of these cloud forests. Moreover, the cloudy, humid environment of these refugial forests appears to exert a strong influence on tree leaf gas exchange and water relations. Because global climate change is predicted to increase regional cloud ceiling levels, more research on cloud impacts on carbon gain and water relations is needed to predict future impacts on these relict forests.     

Differential summer water use by Pinus edulis and Juniperus osteosperma reflects contrasting hydraulic characteristics

Previous studies of pinyon-juniper woodlands show that Pinus edulis Engelm. makes better use of soil water from summer precipitation pulses than does co-occurring Juniperus osteosperma (Torr.) Little. To investigate the basis of this difference, we examined seasonal variation in cavitation and hydraulic conductance. Pinus edulis remained isohydric over the growing season. Minimum water potentials never fell below -2.3 MPa, and the extent of xylem cavitation remained near constant during the dry season. In contrast, J. osteosperma was anisohydric, reaching water potentials as low as -6.9 MPa, and experiencing progressively greater xylem cavitation as the dry season progressed despite having more cavitation-resistant xylem than P. edulis. We conducted an irrigation experiment to observe the responses of the study species to a summer pulse of water. Although sap flow increased in both species in response to the 25-mm irrigation pulse, only J. osteosperma responded to the 10-mm pulse. This was inconsistent with the response of P. edulis to light rain events and may have been due to a difference in the distribution of irrigation water and rain water between the under- and between-canopy areas. Whole-plant conductance increased following the 25-mm irrigation in P. edulis but remained constant in J. osteosperma. We hypothesized that this difference was caused, in part, by differential refilling of embolized xylem. Area specific hydraulic conductivity was 66% higher in roots of irrigated P. edulis trees relative to roots of control trees 3 days after the 25-mm irrigation (t = 2.14, P = 0.02, df = 16). There was no change in hydraulic conductivity of the roots of J. osteosperma or in the stems of either species. Our results indicate that the response to an irrigation pulse in P. edulis depended on cavitation avoidance in stems and the reversal of cavitation in roots, resulting in increased whole-plant conductance and water uptake. In contrast, J. osteosperma failed to exploit light summer rain events but was able to extract deep soil water at low water potentials.     

Water use and carbon exchange of red oak- and eastern hemlock-dominated forests in the northeastern USA: implications for ecosystem-level effects of hemlock woolly adelgid

Water use and carbon exchange of a red oak-dominated (Quercus rubra L.) forest and an eastern hemlock-dominated (Tsuga canadensis L.) forest, each located within the Harvard Forest in north-central Massachusetts, were measured for 2 years by the eddy flux method. Water use by the red oak forest reached 4 mm day(-1), compared to a maximum of 2 mm day(-1) by the eastern hemlock forest. Maximal carbon (C) uptake rate was also higher in the red oak forest than in the eastern hemlock forest (about 25 versus 15 micromol m(-2) s(-1)). Sap flux measurements indicated that transpiration of red oak, and also of black birch (Betula lenta L.), which frequently replaces eastern hemlock killed by hemlock woolly adelgid (Adelges tsugae Annand.), were almost twice that of eastern hemlock. Despite the difference between species in maximum summertime C assimilation rate, annual C storage of the eastern hemlock forest almost equaled that of the red oak forest because of net C uptake by eastern hemlock during unusually warm fall and spring weather, and a near-zero C balance during the winter. Thus, the effect on C storage of replacing eastern hemlock forest with a forest dominated by deciduous species is unclear. Carbon storage by eastern hemlock forests during fall, winter and spring is likely to increase in the event of climate warming, although this may be offset by C loss during hotter summers. Our results indicate that, although forest water use will decrease immediately following eastern hemlock mortality due to the hemlock woolly adelgid, the replacement of eastern hemlock by deciduous species such as red oak will likely increase summertime water use over current rates in areas where hemlock is a major forest species.     

镶黄旗黄花山天然杜松林群落结构与种群分布格局研究

在内蒙古自治区锡林郭勒盟镶黄旗黄花山天然杜松林内,设置4个有代表性的标准样地,应用相邻格子法获得野外调查数据,分析天然杜松林群落结构及乔木层和灌木层优势种种群空间分布格局。结果表明:1)天然杜松林群落内,杜松地径较小,个体高峰值分布范围较为集中,径级结构变化明显;旱榆多以幼苗的形式存在,地径范围在2.5cm以下。群落存在明显的径阶缺陷,属于不稳定型。2)乔木层高度级较低,而灌木层高度级较好。3)乔木层杜松种群主要以聚集分布格局为主,伴有均匀分布格局,旱榆种群则为聚集分布格局;灌木层优势种群小叶鼠李和土庄绣线菊的空间分布格局都为聚集分布。群落处于不稳定的恢复发展阶段,人为和自然干扰以及种群生物学特性共同造成这种空间分布格局。     

Influence of irrigation and fertilization on transpiration and hydraulic properties of Populus deltoides

Long-term hydraulic acclimation to resource availability was explored in 3-year-old Populus deltoides Bartr. ex Marsh. clones by examining transpiration, leaf-specific hydraulic conductance (G(L)), canopy stomatal conductance (G(S)) and leaf to sapwood area ratio (A(L):A(S)) in response to irrigation (13 and 551 mm year(-1) in addition to ambient precipitation) and fertilization (0 and 120 kg N ha(-1) year(-1)). Sap flow was measured continuously over one growing season with thermal dissipation probes. Fertilization had a greater effect on growth and hydraulic properties than irrigation, and fertilization effects were independent of irrigation treatment. Transpiration on a ground area basis (E) ranged between 0.3 and 1.8 mm day(-1), and increased 66% and 90% in response to irrigation and fertilization, respectively. Increases in G(L), G(S) at a reference vapor pressure deficit of 1 kPa, and transpiration per unit leaf area in response to increases in resource availability were associated with reductions in A(L):A(S) and consequently a minimal change in the water potential gradient from soil to leaf. Irrigation and fertilization increased leaf area index similarly, from an average 1.16 in control stands to 1.45, but sapwood area was increased from 4.0 to 6.3 m(2) ha(-1) by irrigation and from 3.7 to 6.7 m(2) ha(-1) by fertilization. The balance between leaf area and sapwood area was important in understanding long-term hydraulic acclimation to resource availability and mechanisms controlling maximum productivity in Populus deltoides.     

Managing forests for wood yield and carbon storage: a theoretical study

Which forest management regimes best achieve the dual objectives of high sustained timber yield and high carbon storage, including the carbon stored in soil and wood products? A mechanistic forest ecosystem simulator, which couples carbon, nitrogen and water (Edinburgh Forest Model), was calibrated to mimic the growth of a pine plantation in a Scottish climate. The model was then run to equilibrium (1) as an undisturbed forest, (2) removing 2.5, 10, 20 or 40% of the woody biomass each year (3) removing 50% of the woody biomass every 20 years, and (4) clear-felling and replanting every 60 years as in conventional plantations in this climate. More carbon was stored in the undisturbed forest (35.2 kg C m(-2)) than in any regime in which wood was harvested. Plantation management gave moderate carbon storage (14.3 kg C m(-2)) and timber yield (15.6 m(3) ha(-1) year(-1)). Notably, annual removal of 10 or 20% of woody biomass per year gave both a high timber yield (25 m(3) ha(-1) year(-1)) and high carbon storage (20 to 24 kg C m(-2)). The efficiency of the latter regimes could be attributed (in the model) to high light interception and net primary productivity, but less evapotranspiration and summer water stress than in the undisturbed forest, high litter input to the soil giving high soil carbon and N(2) fixation, low maintenance respiration and low N leaching owing to soil mineral pool depletion. We conclude that there is no simple inverse relationship between the amount of timber harvested from a forest and the amount of carbon stored. Management regimes that maintain a continuous canopy cover and mimic, to some extent, regular natural forest disturbance are likely to achieve the best combination of high wood yield and carbon storage.