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.     

Botanical determinants of foliage clumping and light interception in two-year-old coppice poplar canopies: assessment from 3-D plant mockups

Botanical parameters (e.g., shoot and branch inclination, petiole length, leaf phyllotaxy, size and shape) that influence light interception and foliage clumping in dense two-year-old monoclonal poplar (Populus spp.) coppice crops were analysed with a three-dimensional simulation model. Crop LAI varied from 1 to 2 for clone Ghoy and from 2.5 to 7.4 for clone Trichobel from May to September. Canopies were strongly clumped, with a clumping index (μ) about 0.5. Canopy light transmittance (τ) varied from 0.59 in May to 0.41 in September for clone Ghoy and from 0.42 to 0.08 for clone Trichobel, and was strongly associated with LAI. The overall effect of a simulated shift in botanical parameters was relatively small and resulted in limited changes in μ and τ by ± 0.05 and ±0.1, respectively. Petiole length had the most notable effect on μ and τ, while the other parameters were less effective. However, biomass cost analyses showed that actual petiole length optimised the efficiency of biomass investment into light capture.     

Drought resistance of two hybrid Populus clones grown in a large-scale plantation

Poplar hybrids were grown with irrigation in a large-scale plantation to investigate the mechanisms underlying clonal differences in drought resistance. Beginning in spring 1992, Populus trichocarpa x P. deltoides (TD) and P. deltoides x P. nigra (DN) cuttings received 46, 76, or 137 cm year(-1) of irrigation to supplement the 18-20 cm of annual precipitation, and all trees received the same fertilization regime. Stem volume, assessed as the square of stem diameter at breast height times tree height (D(2)H), and water relations of the trees were studied from the end of their second growing season until the end of their fifth growing season. By the end of the second growing season, stem volume of Clone TD was 40-146% larger than that of Clone DN, but stem volume growth was independent of irrigation in excess of 46 cm year(-1) in both clones. During the third growing season, stem volume growth of both clones was limited by both the 46- and 76-cm irrigation treatments, so that by the end of the third growing season trees in the 46-cm irrigation treatment were only half the size of trees in the 137-cm irrigation treatment. These treatment differences were maintained through the fifth growing season. Although stem volumes of Clone TD trees in the 76- and 137-cm irrigation treatments were larger than the corresponding values for Clone DN trees at the end of the third growing season (1994), these clonal differences gradually decreased in subsequent years and were not detectable after 5 years, because stem volume relative growth rate of Clone DN was greater than that of Clone TD in all treatments. Although both clones exhibited similar predawn leaf water potentials, Clone DN typically maintained higher midday leaf water potentials, suggesting better stomatal control of water loss. Clonal and treatment differences in osmotic potential at full turgor were minimal and could not explain the clonal differences in drought resistance. Root density and root density to stem volume ratio increased more in response to moderate drought in Clone DN than in Clone TD, resulting in enhanced drought resistance (high stem volume growth rate under moderate drought conditions) and an increased capacity to withdraw water from the soil. We conclude that the greater drought resistance of Clone DN compared with Clone TD was the result of the maintenance of a more favorable water balance by stomatal regulation and greater carbon allocation to roots during the early stages of drought. However, the low root density to stem volume ratio in Clone DN growing in the 46-cm irrigation treatment suggests that severe water limitation restricted the preferential allocation of carbon to belowground tissues, so that both root and shoot growth were constrained by severe drought.     

Hydraulic adjustments underlying drought resistance of Pinus halepensis

Drought-induced tree mortality has increased over the last decades in forests around the globe. Our objective was to investigate under controlled conditions the hydraulic adjustments underlying the observed ability of Pinus halepensis to survive seasonal drought under semi-arid conditions. One hundred 18-month saplings were exposed in the greenhouse to 10 different drought treatments, simulating combinations of intensities (fraction of water supply relative to control) and durations (period with no water supply) for 30 weeks. Stomata closed at a leaf water potential (Ψ(l)) of -2.8 MPa, suggesting isohydric stomatal regulation. In trees under extreme drought treatments, stomatal closure reduced CO(2) uptake to -1 μmol m(-2) s(-1), indicating the development of carbon starvation. A narrow hydraulic safety margin of 0.3 MPa (from stomatal closure to 50% loss of hydraulic conductivity) was observed, indicating a strategy of maximization of CO2 uptake in trees otherwise adapted to water stress. A differential effect of drought intensity and duration was observed, and was explained by a strong dependence of the water stress effect on the ratio of transpiration to evapotranspiration T/ET and the larger partitioning to transpiration associated with larger irrigation doses. Under intense or prolonged drought, the root system became the main target for biomass accumulation, taking up to 100% of the added biomass, while the stem tissue biomass decreased, associated with up to 60% reduction in xylem volume.     

How does drought tolerance compare between two improved hybrids of balsam poplar and an unimproved native species?

Poplars are one of the woody plants that are very sensitive to water stress, which may reduce the productivity of fast-growing plantations. Poplars can exhibit several drought tolerance strategies that may impact productivity differently. Trees from two improved hybrids, Populus balsamifera?×?Populus trichocarpa Torr. & Gray (clone B?×?T) and P. balsamifera?×?Populus maximowiczii A. Henry (clone B?×?M), having P. balsamifera L. as a parent and trees from native and unimproved P. balsamifera were subjected to a 1-month drying cycle in a growth chamber and then rewatered. The unimproved and native B clone maintained higher stomatal conductance (g(s)) than the hybrids, and high photosynthetic activity and transpiration, even when soil water content was nearly zero. As a result, both instantaneous water use efficiency (WUE(i)) and leaf carbon isotope composition (δ(13)C) indicated that this clone was less affected by drought than both hybrids at maximal drought stress. However, this clone shed its leaves when the drought threshold was exceeded, which implied a greater loss of productivity. The B?×?M hybrid showed a relatively conservative response to water stress, with the greatest decrease in transpiring versus absorbing surface (total leaf area to root biomass ratio). This clone was also the only one to develop new leaves after rewatering, and its total biomass production was not significantly decreased by drought. Among the two hybrids, clone B?×?T was the most vigorous, with the greatest transpiration (E(i)) and net CO(2) assimilation (A) rates, allowing for high biomass production. However, it had a more risky strategy under drought conditions by keeping its stomata open and high E(i) rates under moderate drought, resulting in a lower recovery rate after rewatering. The opposite drought response strategies of the two hybrids were reflected by clone B?×?T having lower WUE(i) values than clone B?×?M at maximal drought, with a very low Ψ(min) value of -3.2 MPa, despite closed stomata and stopped photosynthetic activity. Positive linear relationships between A and g(s) for the three hybrids indicated strong stomatal control of photosynthesis. Moreover, the three poplar clones showed anisohydric behaviour for stomatal control and their use under long-term drought should be of interest, especially the B?×?M clone.     

Drought tolerance of clonal Malus determined from measurements of stomatal conductance and leaf water potential

We examined tolerance to soil drying in clonally propagated apple (Malus domestica Borkh.) rootstocks used to control shoot growth of grafted scions. We measured leaf conductance to water vapor (g(L)) and leaf water potential (Psi(L)) in a range of potted, greenhouse-grown rootstocks (M9, M26, M27, MM111, AR69-7, AR295-6, AR360-19, AR486-1 and AR628-2) as the water supply was gradually reduced. Irrespective of the amount of available water, rootstocks that promoted scion shoot growth (M26 and MM111) generally had higher g(L) and more negative Psi(L) than rootstocks that restricted scion shoot growth (M27 and M9). After about 37 days of reduced water supply, there were significant decreases in g(L) and Psi(L) in all rootstocks compared with well-watered controls. In all treatments, the slope of the relationship between log (g(L)) and Psi(L) was positive, except for rootstocks AR295-6, AR628-2 and AR486-1 in the severe-drought treatment, where the drought-induced change in the relationship suggests that rapid stomatal closure occurred when leaf water potentials fell below -2.0 MPa. This drought response was associated with increased root biomass production. Rootstock M26 showed little stomatal closure even when its water potential fell below -2.0 MPa, and there was no effect of drought on root biomass production. We conclude that differences among rootstocks in the way that g(L) and Psi(L) respond to drought reflect differences in the mechanisms whereby they tolerate soil drying. We suggest that these differences are related to differences among the rootstocks in their ability to control shoot growth.     

Variation in drought resistance, drought acclimation and water conservation in four willow cultivars used for biomass production

Growth and water-use parameters of four willow (Salix spp.) clones grown in a moderate drought regime or with ample water supply were determined to characterize their water-use efficiency, drought resistance and capacity for drought acclimation. At the end of the 10-week, outdoor pot experiment, clonal differences were observed in: (1) water-use efficiency of aboveground biomass production (WUE); (2) resistance to xylem cavitation; and (3) stomatal conductance to leaf-specific, whole-plant hydraulic conductance ratio (g(st)/K(P); an indicator of water balance). Across clones and regimes, WUE was positively correlated with the assimilation rate to stomatal conductance ratio (A/g(st)), a measure of instantaneous water-use efficiency. Both of these water-use efficiency indicators were generally higher in drought-treated trees compared with well-watered trees. However, the between-treatment differences in (shoot-based) WUE were smaller than expected, considering the differences in A/g(st) for two of the clones, possibly because plants reallocated dry mass from shoots to roots when subject to drought. Higher root hydraulic conductance to shoot hydraulic conductance ratios (K(R)/K(S)) during drought supports this hypothesis. The same clones were also the most sensitive to xylem cavitation and, accordingly, showed the strongest reduction in g(st)/K(P) in response to drought. Drought acclimation was manifested in decreased g(st), g(st)/K(P), osmotic potential and leaf area to vessel internal cross-sectional area ratio, and increased K(R), K(P) and WUE. Increased resistance to stem xylem cavitation in response to drought was observed in only one clone. It is concluded that WUE and drought resistance traits are inter-linked and that both may be enhanced by selection and breeding.     

Spatial distribution of leaf morphological and physiological characteristics in relation to local radiation regime within the canopies of 3-year-old Populus clones in coppice culture

Spatial distributions of leaf characteristics relevant to photosynthesis were compared within high-density coppice canopies of Populus spp. of contrasting genetic origin. We studied three clones representative of the range in growth potential, leaf morphology, coppice and canopy structure: Clone Hoogvorst (Hoo) (Populus trichocarpa Torr. & Gray x Populus deltoides Bartr. & Marsh), Clone Fritzi Pauley (Fri) (Populus trichocarpa Torr. & Gray) and Clone Wolterson (Wol) (Populus nigra L.). Leaf area index ranged from 2.7 (Fri and Wol) to 3.8 (Hoo). The clones exhibited large vertical variation in leaf area density (0.02-1.42 m2 m-3). Leaf dry mass per unit leaf area (DM(A)) increased with increasing light in Clones Hoo and Fri, from about 56 g m-2 at the bottom of the canopy to 162 g m-2 at the top. In Clone Wol, DM(A) varied only from 65 to 100 g m-2, with no consistent relationship with respect to light. Conversely, nitrogen concentration on a mass basis was nearly constant (around 1.3-2.1%) within the canopies of Clones Hoo and Fri, but increased strongly with light in Clone Wol, from 1.4% at the bottom of the canopy to 4.1% at the top. As a result, nitrogen per unit leaf area (N(A)) increased with light in the canopies of all clones, from 0.9 g m-2 at the bottom to 2.9 g m-2 at the top. Although a single linear relationship described the dependence of maximum carboxylation rate (17-93 micromol CO2 m-2 s-1) or electron transport capacity (45-186 micromol electrons m-2 s-1) on N(A), for all clones, Clone Wol differed from Clones Hoo and Fri by exhibiting a higher dark respiration rate at low N(A) (1.8 versus 0.8 micromol CO2 m-2 s-1).     

Silver birch and climate change: variable growth and carbon allocation responses to elevated concentrations of carbon dioxide and ozone

We studied the effects of elevated concentrations of carbon dioxide ([CO2]) and ozone ([O3]) on growth, biomass allocation and leaf area of field-grown O3-tolerant (Clone 4) and O3-sensitive clones (Clone 80) of European silver birch (Betula pendula Roth) trees during 1999-2001. Seven-year-old trees of Clones 4 and 80 growing outside in open-top chambers were exposed for 3 years to the following treatments: outside control (OC); chamber control (CC); 2 x ambient [CO2] (EC); 2 x ambient [O3] (EO); and 2 x ambient [CO2] + 2 x ambient [O3] (EC+EO). When the results for the two clones were analyzed together, elevated [CO2] increased tree growth and biomass, but had no effect on biomass allocation. Total leaf area increased and leaf abscission was delayed in response to elevated [CO2]. Elevated [O3] decreased dry mass of roots and branches and mean leaf size and induced earlier leaf abscission in the autumn; otherwise, the effects of elevated [O3] were small across the clones. However, there were significant interactions between elevated [CO2] and elevated [O3]. When results for the clones were analyzed separately, stem diameter, volume growth and total biomass of Clone 80 were increased by elevated [CO2] and the stimulatory effects of elevated [CO2] on stem volume growth and total leaf area increased during the 3-year study. Clone 80 was unaffected by elevated [O3]. In Clone 4, elevated [O3] decreased root and branch biomass by 38 and 29%, respectively, whereas this clone showed few responses to elevated [CO2]. Elevated [CO2] significantly increased total leaf area in Clone 80 only, which may partly explain the smaller growth responses to elevated [CO2] of Clone 4 compared with Clone 80. Although we observed responses to elevated [O3], the responses to the EC+EO and EC treatments were similar, indicating that the trees only responded to elevated [O3] under ambient [CO2] conditions, perhaps reflecting a greater quantity of carbohydrates available for detoxification and repair in elevated [CO2].     

Responses to water stress in two Eucalyptus globulus clones differing in drought tolerance

We evaluated drought resistance mechanisms in a drought-tolerant clone (CN5) and a drought-sensitive clone (ST51) of Eucalyptus globulus Labill. based on the responses to drought of some physiological, biophysical and morphological characteristics of container-grown plants, with particular emphasis on root growth and hydraulic properties. Water loss in excess of that supplied to the containers led to a general decrease in growth and significant reductions in leaf area ratio, specific leaf area and leaf-to-root area ratio. Root hydraulic conductance and leaf-specific hydraulic conductance decreased as water stress became more severe. During the experiment, the drought-resistant CN5 clone maintained higher leaf water status (higher predawn and midday leaf water potentials), sustained a higher growth rate (new leaf area expansion and root growth) and displayed greater carbon allocation to the root system and lower leaf-to-root area ratio than the drought-sensitive ST51 clone. Clone CN5 possessed higher stomatal conductances at moderate stress as well as higher hydraulic conductances than Clone ST51. Differences in the response to drought in root biomass, coupled with changes in hydraulic properties, accounted for the clonal differences in drought tolerance, allowing Clone CN5 to balance transpiration and water absorption during drought treatment and thereby prolong the period of active carbon assimilation.     

Metabolic responses to water deficit in two Eucalyptus globulus clones with contrasting drought sensitivity

We compared the metabolic responses of leaves and roots of two Eucalyptus globulus Labill. clones differing in drought sensitivity to a slowly imposed water deficit. Responses measured included changes in concentrations of soluble and insoluble sugars, proline, total protein and several antioxidant enzymes. In addition to the general decrease in growth caused by water deficit, we observed a decrease in osmotic potential when drought stress became severe. In both clones, the decrease was greater in roots than in leaves, consistent with the observed increases in concentrations of soluble sugars and proline in these organs. In roots of both clones, glutathione reductase activity increased significantly in response to water deficit, suggesting that this enzyme plays a protective role in roots during drought stress by catalyzing the catabolism of reactive oxygen species. Clone CN5 has stress avoidance mechanisms that account for its lower sensitivity to drought compared with Clone ST51.     

Diurnal patterns of water use in Eucalyptus victrix indicate pronounced desiccation-rehydration cycles despite unlimited water supply

Knowledge about nocturnal transpiration (E(night)) of trees is increasing and its impact on regional water and carbon balance has been recognized. Most of this knowledge has been generated in temperate or equatorial regions. Yet, little is known about E(night) and tree water use (Q) in semi-arid regions. We investigated the influence of atmospheric conditions on daytime (Q(day)) and nighttime water transport (Q(night)) of Eucalyptus victrix L.A.S. Johnson & K.D. Hill growing over shallow groundwater (not >1.5 m in depth) in semi-arid tropical Australia. We recorded Q(day) and Q(night) at different tree heights in conjunction with measurements of stomatal conductance (g(s)) and partitioned E(night) from refilling processes. Q of average-sized trees (200-400 mm diameter) was 1000-3000 l month(-1), but increased exponentially with diameter such that large trees (>500 mm diameter) used up to 8000 l month(-1). Q was remarkably stable across seasons. Water flux densities (J(s)) varied significantly at different tree heights during day and night. We show that g(s) remained significantly different from zero and E(night) was always greater than zero due to vapor pressure deficits (D) that remained >1.5 kPa at night throughout the year. Q(night) reached a maximum of 50% of Q(day) and was >0.03 mm h(-1) averaged across seasons. Refilling began during afternoon hours and continued well into the night. Q(night) eventually stabilized and closely tracked D(night). Coupling of Q(night) and D(night) was particularly strong during the wet season (R2?=?0.95). We suggest that these trees have developed the capacity to withstand a pronounced desiccation-rehydration cycle in a semi-arid environment. Such a cycle has important implications for local and regional hydrological budgets of semi-arid landscapes, as large nighttime water fluxes must be included in any accounting.     

Post-fertilization physiology and growth performance of loblolly pine clones

The physiological processes leading to enhanced growth of loblolly pine (Pinus taeda L.) following fertilization are not clearly understood. Part of the debate revolves around the temporal response of net photosynthetic rate (A(n)) to fertilization and whether the A(n) response is always positive. We measured light-saturated photosynthetic rate (A(sat)), dark respiration rate, growth and crown silhouette area in eight clones of loblolly pine before and after nitrogen (N) fertilization (112 kg ha(-1)) to track the initial physiological changes prior to any changes in growth. Overall, there were positive photosynthetic and growth responses to fertilization; however, there were pronounced physiological and growth differences among clones, even among clones with the same parents. Clones 4, 6 and 7 showed large volume growth and A(sat) responses to fertilization. Clone 1 and Clone 8 (a full-sibling of Clone 7) mainly showed a volume growth response, whereas Clone 2 (full-sibling of Clone 1) showed an A(sat) response only. Clone 5 (full-sibling of Clone 6) showed little response to fertilization, whereas Clone 3 (full-sibling of Clone 4) showed a negative A(sat) response. Thus, within-family variation warrants further study to ensure that relatively expensive clonal material is used efficiently.     

Stomatal patchiness in the Mediterranean holm oak (Quercus ilex L.) under water stress in the nursery and in the forest

The evergreen holm oak Quercus ilex L. is the most representative tree in Mediterranean forests. Accurate estimation of the limiting factors of photosynthesis for Q. ilex and the prediction of ecosystem water-use efficiency by mechanistic models can be achieved only by establishing whether this species shows heterogenic stomatal aperture, and, if so, the circumstances in which this occurs. Here, we collected gas-exchange and chlorophyll fluorescence data in Q. ilex leaves from a nursery to measure the effects of stomatal oscillations on PSII quantum yield (Φ(PSII)) under water stress. Stomatal conductance (g(s)) was used as an integrative indicator of the degree of water stress. Images of chlorophyll fluorescence showed heterogeneous Φ(PSII) when g(s) was <50 mmol H(2)O m(-2) s(-1), representative of severe drought and corresponding to a container capacity <45%. Stomatal patchiness was related to a coefficient of variation (CV) of Φ(PSII) values >2.5%. A parallel study in the forest confirmed heterogeneous Φ(PSII) values in leaves in response to declining water availability. Three kinds of Q. ilex individuals were distinguished: those resprouting after a clear-cut (resprouts, R); intact individuals growing in the same clear-cut area as resprouts (controls, C); and intact individuals in a nearby, undisturbed area (forest controls, CF). Patchiness increased in C and CF in response to increasing drought from early May to late July, whereas in R, Φ(PSII) values were maintained as a result of their improved water relations since the pre-existing roots were associated with a smaller aerial biomass. Patchiness was related to a % CV of Φ(PSII) values >4 and associated in the summer with mean g(s) values of 30 mmol H(2)O m(-2) s(-1). Under milder drought in spring, Φ(PSII) patchiness was less strictly related to g(s) variations, pointing to biochemical limitants of photosynthesis. The occurrence of heterogenic photosynthesis caused by patchy stomatal closure in Q. ilex during severe drought should be taken into account in ecosystem modelling in which harsher water stress conditions associated with climate change are predicted.     

Use of a single-tree simulation model to predict effects of ozone and drought on growth of a white fir tree

A physiologically based, single-tree simulation model, TREGRO, was parameterized with existing phenological, allometric, and growth data and used to predict effects of ozone and drought on growth of a 53-year-old white fir (Abies concolor (Gord. & Glend.) Lindl. ex Hildebr.) tree following a 3-year model simulation. Multiple experimental simulations were conducted to assess the individual and interactive effects of ozone (O(3)) exposure and drought on growth of white fir. The effects of O(3) were imposed as reductions in carbon (C) assimilation of 0, 2.5, 5, 10, and 20%. Drought was imposed as 0, 10, 25, and 50% reductions in total annual precipitation. The results of the simulations were compared with the effects of O(3) on white fir seedlings grown in the presence and absence of ozone in open-top chambers and with a field survey of white fir trees subjected to a gradient of O(3). In the O(3) simulations, an O(3)-induced reduction in C assimilation of 2.5% reduced total tree biomass and branch total nonstructural carbohydrate (TNC) content by < 7%. Although quantifiable in simulation experiments, such small reductions would probably not be detectable in the field. Results from both an open-top chamber experiment and a field survey indicated that reductions in C assimilation of white fir growing in elevated O(3) were much greater than 2.5%, but were not statistically different from control values. A simulated O(3) reduction in C assimilation of >/= 10% reduced total tree biomass by 7% and branch TNC by 55%. Results from the field survey indicated that branch elongation was reduced in response to increased O(3) concentration, corroborating the simulated response of reduced C allocation to the branches of white fir. Although simulated reductions in total annual precipitation of >/= 25% reduced final tree biomass, the simulated reductions also reduced O(3) uptake and therefore reduced the O(3) response of white fir. However, a combination of low amounts of O(3) (2.5% reduction in C assimilation) and drought (25% reduction in annual precipitation) synergistically reduced C gain of white fir more than either stress individually. Our simulations predict that moderate drought (no more than a 25% reduction in total annual precipitation) may not ameliorate the response of white fir to O(3) and that moderate amounts of atmospheric O(3) and drought could be more detrimental to white fir than either stress singly.     

Through droughts and hurricanes: Tree mortality,forest structure,and biomass production in a coastal swamp targeted for restoration in the Mississippi River Deltaic Plain

Coastal swamps are among the rapidly vanishing wetland habitats in Louisiana due to accelerated sea-level rise and hydrological alterations that alter the natural flooding regime. In particular, the swamp forests of Lake Maurepas, Louisiana, have degraded considerably, and research regarding their condition might suggest approaches for their restoration. We measured forest structure, species composition, tree mortality, annual aboveground net primary production (ANPP) of woody species, and aboveground biomass allocation to leaf litter and wood, and soil strength at forty study plots within the Lake Maurepas basin over 5 years to evaluate the current condition of this coastal forested wetland. Local measures of salinity and regional measures of flooding were used to predict ANPP and aboveground biomass allocation. The 5-year study period included an intense drought as well as years characterized by hurricane-induced flooding. The forty study plots could be divided into four distinct habitat clusters based on standing biomass, structural variables, and salinity. The majority of the plots were co-dominated by Taxodium distichum and Nyssa aquatica. Acer rubrum var. drummondii and Fraxinus pennsylvanica were common mid-story species throughout the western and southern parts of the study area, while Salix nigra, Morella cerifera, and Triadica sebiferum were more important at the more degraded plots in the eastern part of the basin. Annual mean soil salinity reached unprecedented level (2–5 psu) during the drought and cumulative tree mortality reached up to 85% in areas characterized by frequent saltwater intrusions. The ANPP was higher during the drought period in 2000–2001 than during subsequent years, and was dominated by T. distichum. At most sampling plots, litter production exceeded wood production annually. A negative correlation between aboveground biomass allocation to litter and flooding indicated that biomass allocation shifted from litter toward wood during wet years. Overall, the majority of the plots sampled produced less than 400 g m⁻² yr⁻¹ of aboveground biomass annually due to the interacting negative effects of saltwater intrusion and prolonged flooding with nutrient-poor water. Reintroduction of Mississippi River water to the Maurepas system has the potential to benefit these swamps greatly by restoring a greater flow of nutrients, sediments, and fresh water through the wetlands. The historically slow (i.e., multi-decadal) process of swamp deterioration was greatly sped by low salinity (i.e., 2–5 psu) saltwater intrusions during a drought in 1999–2000. The majority of the coastal swamps in the Pontchartrain Basin are deteriorating, and most of this swamp area will be lost to open water in the foreseeable future if no restoration action is taken.     

Growth,biomass allocation,and water use efficiency of 31 apple cultivars grown under two water regimes

Plant growth, biomass allocation, carbon isotope composition (δ¹³C), and water use efficiency (WUE) of 31 cultivars of apple (Malus domestica Borkh.) grown under two water regimes were measured. Drought-stressed plants showed significant declines in tree height, trunk diameter, biomass production, and total leaf area, the extent to which depended upon cultivar. Also, gas exchange rates, instantaneous and long-term efficiencies (WUE_I and WUE_L, respectively), and values for δ¹³C differed among cultivars and watering regimes. Variations in WUE_I were mainly due to changes in stomatal conductance (g _s) under drought condition. ‘Qinguan’ and ‘Golden Delicious’ had greater trunk diameter, tree height, and had higher biomass production and WUE_L under drought stress, implying that they are more suitable for arid and semi-arid regions. Moreover, WUE_L was significantly and positively correlated with δ¹³C under two watering regimes, which suggests a potential for evaluating water use efficiency of Malus by measuring carbon isotope composition.     

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.     

Changes in the relationship between tree size and aboveground respiration in field-grown hinoki cypress (Chamaecyparis obtusa) trees over three years

Respiration measurements of aerial parts of 18-year-old hinoki cypress (Chamaecyparis obtusa (Sieb. et Zucc.) Endl.) trees were made under field conditions over three years to study changing relationships with tree age between respiration and phytomass, phytomass increment, and leaf mass. The relationship between annual respiration (r(a)) and phytomass (w(T)) was approximated by a proportional function (r(a) = aw(T)), where the proportional constant (a) decreased year by year. The effect of time on the relationship between annual respiration and phytomass of each sample tree was fitted by a power function. Respiration of the tree suppressed by the canopy decreased year by year, but respiration of the other trees increased slightly with age. The relationship between annual respiration and leaf mass was also approximated by a generalized power function. Excluding the suppressed tree, the relationship between annual respiration (r(a)) and the annual increment of aboveground phytomass (Deltaw(T)) was described by a proportional function (r(a) = 2.27Deltaw(T)), where the proportional constant, 2.27, was independent of sample tree and year, indicating that about 2.3 times of the annual aboveground phytomass increment equivalent was respired annually. For any tree, the time constant relationships between annual respiration and leaf mass and phytomass increment for different-sized trees were similar to the corresponding time continuum relationships. In contrast, the time continuum relationship between annual respiration and phytomass differed from the time constant relationship, indicating that respiration of less active woody tissue contributed significantly to aboveground respiration. Based on the relationship between tree size and annual respiration, annual aboveground stand respiration was estimated to be 25.0, 26.9, and 25.8 Mg(dm) ha(-1) year(-1) for the three consecutive years, respectively, and the corresponding aboveground stand biomass was 60.0, 69.0, and 76.8 Mg(dm) ha(-1).     

Clonal and seasonal differences in leaf osmotic potential and organic solutes of five hybrid poplar clones grown under field conditions

Leaf osmotic potential at full turgor (Psi(pio)) and the major solutes that contribute to osmotic potential were characterized in five hybrid poplar clones of Populus trichocarpa Torr. & Gray x P. deltoides Bartr. (TD) and P. deltoides x P. nigra L. (DN), growing under field conditions at two sites in eastern Washington and Oregon, USA. Trees were drip irrigated with 46, 76 or 137 cm of supplemental irrigation during each growing season. Trees at Wallula, WA, which were in their third growing season in 1994, were sampled twice a year for two years (1994 and 1995), and trees at Boardman, OR, which were in their second growing season in 1994, were sampled once a year for three years (1994-1996). At Wallula, the TD and DN clones exhibited lower predawn leaf water potentials in the 46-cm treatment than in the 137-cm treatment (-1.2 versus -0.7 MPa) during a hot, dry period in July 1994. Clone TD had a lower Psi(pio) than Clone DN (-1.67 versus -1.56 MPa) during the same period and the difference was also evident in 1995 (-1.81 versus -1.72 MPa) when trees were in their fourth growing season. There was also a significant treatment effect on Psi(pio) in Clone TD, with trees in the 46-cm treatment having lower Psi(pio) than trees in the 137-cm treatment in July 1994. At Boardman, Psi(pio) was generally high with no treatment differences during the 1994-96 samplings. The TD clones had significantly lower Psi(pio) than the DN clones in 1994 (-1.44 versus -1.36 MPa) and 1996 (-1.72 versus -1.54 MPa), but there was no difference between clones in 1995 (-1.40 versus -1.43 MPa). In 1995, at Wallula, osmotic adjustment in Clone TD was largely accounted for by an increase in sucrose, which constituted 70% of total organic solutes. Although the total concentration of free primary amino acids in this clone was 28% higher in trees in the 46-cm treatment than in trees in the 137-cm treatment, amino acids constituted only a small fraction of the total solute pool. Sixty-two percent of total solutes were inorganic ions in Clone TD compared to 52% in Clone DN, and potassium was the main ion constituting about 30% of total solutes and 50% of total ions. However, the clonal difference in Psi(pio) was not fully accounted for by the difference in solute concentration. Osmotic potential at full turgor declined over the growing season and with age. We conclude that, because the extent of osmotic adjustment exhibited by these clones was small, other drought resistance mechanisms contributed to the clonal differences in field performance.