Leaf gas exchange of trees in old-growth and young secondary forest stands in Sulawesi, Indonesia

In the tropics, old-growth forests are converted to other land cover types at a high rate and young secondary forest may gain in importance. Information on associated changes in leaf gas exchange and other leaf traits can be valuable for modelling biogeochemical fluxes under altered land-use patterns. We studied in situ photosynthetic parameters and stomatal conductance for water vapour in eight abundant tree species of young secondary forest and eight tree species of natural old-growth forest in Central Sulawesi, Indonesia. In sun leaves, the average maximal stomatal conductance (g _smax) in the secondary forest (SF) species was 2.1 times higher than in the old-growth forest (OGF) species. Species with a high g _smax reduced g _s sharply when vapour pressure deficit of the air increased, whereas species with a low g _smax were much less sensitive to air humidity. For area-based photosynthetic capacity (A _max-area), the SF species had a 2.3 times higher average than the OGF species. For both, g _smax and A _max-area the variation among species was higher in the OGF than in the SF. When all tree species (n=16) are considered, species means of specific leaf area (SLA), leaf N concentration and leaf P concentration were significantly correlated with g _smax and A _max-area. The strong correlation between A _max-area and foliar P (r ²=0.8) is remarkable as the alluvial soils in the study region are rich in nutrients. If the eight OGF species are analysed separately, the only significant correlation was observed between SLA and mass-based A _max; in the SF species strong correlations were found between leaf size and A _max-area and g _smax. These results show that the conversion of old-growth forest to young secondary forest in Sulawesi significantly alters tree leaf gas exchange characteristics and that chemical and structural leaf traits can be used for the prediction of these changes. The best correlations between leaf gas exchange parameters and leaf traits were obtained by different traits in the SF species, the OGF species and the entire pool of studied species.     

Specific leaf area and leaf nitrogen concentration in annual and perennial grass species growing in Mediterranean old-fields

 We evaluated the hypothesis that photosynthetic traits differ between leaves produced at the beginning (May) and the end (November–December) of the rainy season in the canopy of a seasonally dry forest in Panama. Leaves produced at the end of the wet season were predicted to have higher photosynthetic capacities and higher water-use efficiencies than leaves produced during the early rainy season. Such seasonal phenotypic differentiation may be adaptive, since leaves produced immediately preceding the dry season are likely to experience greater light availability during their lifetime due to reduced cloud cover during the dry season. We used a construction crane for access to the upper canopy and sampled 1- to 2-month-old leaves marked in monthly censuses for six common tree species with various ecological habits and leaf phenologies. Photosynthetic capacity was quantified as light- and CO₂-saturated oxygen evolution rates with a leaf-disk oxygen electrode in the laboratory (O_2max) and as light-saturated CO₂ assimilation rates of intact leaves under ambient CO₂ (A_max). In four species, pre-dry season leaves had significantly higher leaf mass per unit area. In these four species, O_2max and A_max per unit area and maximum stomatal conductances were significantly greater in pre-dry season leaves than in early wet season leaves. In two species, A_max for a given stomatal conductance was greater in pre-dry season leaves than in early wet season leaves, suggesting a higher photosynthetic water-use efficiency in the former. Photosynthetic capacity per unit mass was not significantly different between seasons of leaf production in any species. In both early wet season and pre-dry season leaves, mean photosynthetic capacity per unit mass was positively correlated with nitrogen content per unit mass both within and among species. Seasonal phenotypic differentiation observed in canopy tree species is achieved through changes in leaf mass per unit area and increased maximum stomatal conductance rather than by changes in nitrogen allocation patterns. Received: 7 March 1996 / Accepted: 1 August 1996     

Late-stage canopy tree species with extremely low δ13C and high stomatal sensitivity to seasonal soil drought in the tropical rainforest of French Guiana

We assessed the daily time‐courses of CO₂ assimilation rate (A), leaf transpiration rate (E), stomatal conductance for water vapour (g_s), leaf water potential ( Ψ _w) and tree transpiration in a wet and a dry season for three late‐stage canopy rainforest tree species in French Guiana differing in leaf carbon isotope composition ( δ ¹³C). The lower sunlit leaf δ ¹³C values found in Virola surinamensis ( ? 29·9‰) and in Diplotropis purpurea ( ? 30·9‰), two light‐demanding species, as compared to Eperua falcata ( ? 28·6‰), a shade‐semi‐tolerant species, were clearly associated with higher maximum g_s values of sunlit leaves in the two former species. These two species were also characterized by a high sensitivity of g_s, sap flow density (Ju) and canopy conductance (g_c) to seasonal soil drought, allowing maintenance of high midday Ψ _w values in the dry season. The data for Diplotropis provided an original picture of increasing midday Ψ _w with increasing soil drought. In Virola, stomata were extremely sensitive to seasonal soil drought, leading to a dramatic decrease in leaf and tree transpiration in the dry season, whereas midday Ψ _w remained close to ? 0·3 MPa. The mechanisms underlying such an extremely high sensitivity of stomata to soil drought remain unknown. In Eperua, g_s of sunlit leaves was non‐responsive to seasonal drought, whereas Ju and g_c were lower in the dry season. This suggests a higher stomatal sensitivity to seasonal drought in shaded leaves than in sunlit ones in this species.     

Canopy gradients in leaf intercellular CO2 mole fractions revisited: interactions between leaf irradiance and water stress need consideration

Intercellular CO₂ mole fractions (C_i) are lower in the upper canopy relative to the lower canopy leaves. This canopy gradient in C_i has been associated with enhanced rates of carbon assimilation at high light, and concomitant greater draw‐downs in C_i. However, increases in irradiance in the canopy are generally also associated with decreases in leaf water availability. Thus, stress effects on photosynthesis rates (A) and stomatal conductance (G), may provide a further explanation for the observed C_i gradients. To test the hypotheses of the sources of canopy variation in C_i, and quantitatively assess the influence of within‐canopy differences in stomatal regulation on A, the seasonal and diurnal variation in G was studied in relation to seasonal average daily integrated quantum flux density (Q_int) in tall shade‐intolerant Populus tremula L. trees. Daily time‐courses of A were simulated using the photosynthesis model of Farquhar et al. (Planta 149, 78–90, 1980). Stable carbon isotope composition of a leaf carbon fraction with rapid turnover rate was used to estimate canopy gradient in C_i during the simulations. Daily maximum G (G_max) consistently increased with increasing Q_int. However, canopy differences in G_max decreased as soil water availability became limiting during the season. In water‐stressed leaves, there were strong mid‐day decreases in G that were poorly associated with vapour pressure deficits between the leaf and atmosphere, and the magnitude of the mid‐day decreases in G occasionally interacted with long‐term leaf light environment. Simulations indicated that the percentage of carbon lost due to mid‐day stomatal closure was of the order of 5–10%, and seasonal water stress increased this percentage up to 20%. The percentage of carbon lost due to stomatal closure increased with increasing Q_int. Canopy differences in light environment resulted in a gradient of daily average C_i of approximately 20 µmol mol^?1. The canopy variation in seasonal and diurnal reductions in G led to a C_i gradient of approximately 100 µmol mol^?1, and the actual canopy C_i gradient was of the same magnitude according to leaf carbon isotope composition. This study demonstrates that stress effects influence C_i more strongly than within‐canopy light gradients, and also that leaves acclimated to different irradiance and water stress conditions may regulate water use largely independent of foliar photosynthetic potentials.     

Diurnal patterns of carbon dioxide,water vapour,and energy fluxes in pineapple [<Emphasis Type="Italic">Ananas comosus</Emphasis> (L.) Merr. cv. Red Spanish] field using eddy covariance

We analyzed the eddy covariance measurements of momentum, mass, and energy taken daily throughout five consecutive seasonal courses (i.e. 840 d after planting) of a pineapple [Ananas comosus (L.) Merr. cv. Red Spanish] field growing in the Orinoco lowlands. This field provides an opportunity for micrometeorological studies because of the flat and windy site; the seasonal weather including ENSO effects and the Crassulacean Acid Metabolism (CAM) physiology of the crop were additional attributes. Soil CO₂ flux was quantified and added to the net ecosystem exchange in order to obtain the canopy flux (F_C). The canopy CO₂ flux partially followed the four phases of CAM sensu Osmond (1978). The daily pattern of gaseous exchange in pineapple showed a continuum spectrum in which a major proportion of CO₂ uptake occurring during the daytime was common and in which the CAM expression was related to day and nocturnal CO₂ uptake. However, the benefits of CO₂ uptake at low water cost were constrained by the limited nocturnal CO₂ uptake. Seasonal and ontogenetic changes affected the energy exchange as well as the partitioning of available energy into sensible (Q_H) and latent (Q_LE) heat. When the hourly net radiation (Q_Rn) reached its maximum value, latent heat flux (Q_LE) to available energy throughout the vegetative and reproductive stages was 0.65, 0.05, 0.30, 0.11, and 0.33 for the 1997 wet season, 1997/98 dry season, 1998 wet season, 1998/99 dry season, and 1999 wet season, respectively. Throughout the growth period, we found the pivotal role of surface conductance (g _S) in both Q_LE and F_C. Furthermore, the canopy responded to environmental changes. During the wet seasons the g _S was strongly influenced by humidity mole fraction deficit and was usually lower than aerodynamic conductance, whereas during the dry seasons, soil water deficit limited evapotranspiration and production rates. For the fully canopy cover, the hourly trend of marginal water cost of pineapple carbon gain in the dry seasons indicated that g _S became sufficiently efficient to reduce the amount of water transported per unit of carbon gain. In the wet season, the coupling of CO₂ uptake and stomatal conductance was more effective in maintaining a higher proportionality between Q_LE and g _S.     

Wet- vs. Dry-Season Transpiration in an Amazonian Rain Forest Palm Iriartea deltoidea

The wet and dry seasons in tropical rain forests can differ in precipitation, soil moisture and irradiance more significantly than often assumed. This could potentially affect the water relations of many tree species that may exhibit either increased transpiration in the dry season as a response to the increased irradiance or decreased transpiration as a result of decreases in soil moisture and increases in atmospheric vapor pressure deficit (VPD). Atmospheric data, soil moisture data and sap fluxes in Iriartea deltoidea palms were measured in eastern Ecuador during the wet and dry seasons. There were no differences between total daily sap fluxes in I. deltoidea palms during the wet and dry seasons; however, evaporative demand was significantly higher in the dry season and therefore, transpiration was more restricted by stomatal closure during the dry season than the wet season. This is likely the result of larger atmospheric VPD during the dry season compared with the wet season and possibly the result of reduced soil moisture availability. Additionally, based on published tree abundances in this area, measured sap fluxes in I. deltoidea were scaled up to the hectare level. Transpiration from I. deltoidea palms was estimated to be around 0.03 mm/d, which could represent about 1 percent of total transpiration in this area of the Amazon rain forest. If climate change predictions for more lengthy tropical dry periods are realized, greater stomatal control of dry-season sap flux has the potential to become even more prevalent in tropical species.     

Photosynthetic responses of Miconia species to canopy openings in a lowland tropical rainforest

We examined the photosynthetic acclimation of three tropical species of Miconia to canopy openings in a Costa Rican rainforest. The response of photosynthesis to canopy opening was very similar in Miconia affinis, M. gracilis, and M. nervosa, despite differences in growth form (trees and shrubs) and local distributions of plants (understory and gap). Four months after the canopy was opened by a treefall, photosynthetic capacity in all three species had approximately doubled from closed canopy levels. There were no obvious signs of high light damage after treefall but acclimation to the gap environment was not immediate. Two weeks after treefall, A_max, stomatal conductance, apprarent quantum efficiency, and dark respiration rates had not changed significantly from understory values. The production of new leaves appears to be an important component of light acclimation in these species. The only variables to differ significantly among species were stomatal conductance at A_max and the light level at which assimilation was saturated. M. affinis had a higher stomatal conductance which may reduce its water use efficiency in gap environments. Photosynthesis in the more shade-tolerant M. gracilis saturated at lower light levels than in the other two species. Individual plant light environments were assessed after treefall with canopy photography but they explained only a small fraction of plant variation in most measures of photosynthesis and growth. In conclusion, we speculate that species differences in local distribution and in light requirements for reproduction may be more strongly related to species differences in carbon allocation than in carbon assimilation.     

Canopy‐scale relationships between stomatal conductance and photosynthesis in irrigated rice

Modeling stomatal behavior is critical in research on land–atmosphere interactions and climate change. The most common model uses an existing relationship between photosynthesis and stomatal conductance. However, its parameters have been determined using infrequent and leaf‐scale gas‐exchange measurements and may not be representative of the whole canopy in time and space. In this study, we used a top‐down approach based on a double‐source canopy model and eddy flux measurements throughout the growing season. Using this approach, we quantified the canopy‐scale relationship between gross photosynthesis and stomatal conductance for 3 years and their relationships with leaf nitrogen content throughout each growing season above a paddy rice canopy in Japan. The canopy‐averaged stomatal conductance (g_sc) increased with increasing gross photosynthesis per unit green leaf area (A_g), as was the case with leaf‐scale measurements, and 41–90% of its variation was explained by variations in A_g adjusted to account for the leaf‐to‐air vapor‐pressure deficit and CO₂ concentration using the Leuning model. The slope (m) in this model (g_sc versus the adjusted A_g) was almost constant within a 15‐day period, but changed seasonally. The m values determined using an ensemble dataset for two mid‐growing‐season 15‐day periods were 30.8 (SE = 0.5), 29.9 (SE = 0.7), and 29.9 (SE = 0.6) in 2004, 2005, and 2006, respectively; the overall mid‐season value was 30.3 and did not greatly differ among the 3 years. However, m appeared to be higher during the early and late growing seasons. The ontogenic changes in leaf nitrogen content strongly affected A_g and thus g_sc. In addition, we have discussed the agronomic impacts of the interactions between leaf nitrogen content and g_sc. Despite limitations in the observations and modeling, our canopy‐scale results emphasize the importance of continuous, season‐long estimates of stomatal model parameters for crops using top‐down approaches.     

A comparison of sap flux and water relations of leaves of various isolated trees with special reference to foundation movement in clay soil

Diurnal variation in sap flux (S) through stems of six trees, two each of Ulmus procera SALISB., Melaleuca styphelioides SM. and Prunus cerasifera EHRH. ‘Nigra’ (referred to hereafter by their generic names), were estimated from measurements of heat pulse velocities. Leaf water potential (ψ), stomatal conductance (g _s ) and transpiration from leaves (T) of all replicate trees were measured at 1300–1500h, once during the summer. On two separate occasions measurements were made of S, ψ, (g _s ) and T for one each of Ulmus and Melaleuca trees to study diurnal variations in these parameters. A 12×12 m² area around each tree was kept covered to simulate the condition of trees growing on pavements adjacent to residential properties. Sap flux for these tree species was in the order Melaleuca>Ulmus>Prunus. It is suggested that the smaller canopy and sapwood area in Prunus compared to the other two species is responsible for lower water potential and lower transpiration rate than the other species. Detailed analysis of the diurnal variation in sap flux and water relation of leaves of Melaleuca and Ulmus indicated sap flux of Melaleuca to be greater than that of Ulmus at the same transpiration rate per unit leaf area although the sapwood area of the two species was marginally different. This may have been due either to the difference in canopy conductance or in leaf area between the two species. With the assumption that sap flux closely resembles the rate of soil water extraction for both species, results indicate that Melaleuca is likely to extract soil water at a higher rate than Ulmus and hence is capable of causing greater shrinkage and soil movement than Ulmus.     

Gas exchange and resource-use efficiency of Leymus cinereus (Poaceae): diurnal and seasonal responses to naturally declining soil moisture

We examined factors that limit diurnal and seasonal photosynthesis in Leymus cinereus, a robust tussock grass from shrub-steppes of western North America. Data from plants in a natural stand and in experimental field plots indicate that this bunchgrass has 1) a high photosynthetic capacity, 2) high leaf nitrogen content and high nitrogen-use efficiency, 3) a steep leaf-to-air diffusion gradient for carbon dioxide, which enhances intrinsic water-use efficiency, and 4) photosynthetic tissues that tolerate severe water stress and recover quickly from moderate water stress. Midday depressions of CO₂ assimilation (A) and stomatal conductance were slight in plants with plentiful water, but marked in plants subject to moderate water stress. Midday stomatal closure in moderately stressed plants reduced intercellular carbon dioxide concentration (c_i) by ≈40 μl liter^-1. The maximum rate of A achieved during the day for severely stressed plants (predawn water potential = -4 MPa) was one-third and daily carbon gain per unit leaf area was about one-fourth that of well-watered plants. For plants in the natural stand, CO₂-saturated photosynthesis declined almost linearly with decreasing soil water availability over the growing season, whereas there was little effect on A at CO₂ ambient levels or on carboxylation efficiency until predawn water potentials reached -1.8 MPa. Nitrogen-use efficiency declined with diminishing soil moisture, but there was no seasonal change in stomatal limitation or instantaneous water-use efficiency as estimated from A vs. c_i curves at optimal leaf temperature and moderate atmospheric evaporative demand. Thus, reduced stomatal conductance in response to increased evaporative demand may increase stomatal limitation diumally, but over the growing season, stomatal limitation estimated from A vs. c_i curves is relatively constant because maximum stomatal conductance is closely tuned to the CO₂ assimilatory capacity of the mesophyll.     

Responses of sap flux and stomatal conductance of Pinus taeda L. trees to stepwise reductions in leaf area

Herbivory or artificial foliage removal has been shown to affect gas exchange and canopy water relations. In this study, canopy architecture and water relations in response to progressive defoliation were examined in a stand of 8-year-old loblolly pine (Pinus taeda L.) trees, a shade-intolerant, pioneer species common in the south-eastern USA. Sap flux was measured with constant heat sap flow gauges in order to estimate canopy stomatal conductance (Gs) while foliage in the 6 m high stand was harvested in 1 m increments from the bottom up. Leaf-level stomatal conductance and water potential data were also collected. Profiles of silhouette area ratio and specific leaf area showed no trends with crown height, reflecting an open canopy (leaf area index = 1.55). Therefore, short-term changes in Gs with foliage removal were attributed to hydraulic effects rather than influences of changes in mean microclimate conditions on Gs of remaining foliage. A large increase in Gs was observed during the 6 h pruning period which fully compensated for the reductions in foliage area down to 45%. Canopy stomatal conductance and whole plant liquid phase conductance as calculated from sap flux were both influenced by the rate of growth as indicated by the annual basal area increment.     

Temporal patterns of water flux in trees and lianas in a Panamanian moist forest

Using constant heat sap flow sensors, xylem water fluxes in ten tree species and two liana species were monitored for 5–10 days during the beginning of the wet season in May, 1993. For a subset of the trees, a branch was also monitored at the top of the crown for 5 days. Xylem flux (J _S) was related diurnally in all plants to vapor pressure deficit (D) measured within the upper-third of the canopy, and to incoming shortwave radiation R _S above the canopy. Cross-correlation analysis was used to estimate time lags between diurnal patterns of J _S and D or R _S, and between J _S in stems and branches. The maximum correlation coefficient from cross-correlation of J _S with R _S (range=0.57–0.92) was often higher than the maximum of J _S with D (range=0.43–0.89), indicating that diurnal J _S was more dependent on R _S than D. Time lags (lag corresponding to maximum correlation) of J _S at stem-base with D was shorter (0–45 min) than with radiation (5–115 min), highly variable within a species, and uncorrelated to the height or exposure of tree crowns or liana in the canopy. On a stand level, not accounting for the diel lag between stem sap flux and canopy flux resulted in errors in estimated canopy transpiration of up to 30%. Received: 19 October 1998 / Accepted: 8 June 1999     

Gender specific patterns of carbon uptake and water use in a dominant riparian tree species exposed to a warming climate

Air temperatures in the arid western United States are predicted to increase over the next century. These increases will likely impact the distribution of plant species, particularly dioecious species that show a spatial segregation of the sexes across broad resource gradients. On the basis of spatial segregation patterns, we hypothesized that temperature increases will have a greater negative impact on female plants compared with co‐occurring male plants of dioecious species. This hypothesis was tested by examining the whole‐plant carbon and water relations of 10‐year‐old female (n = 18) and male (n = 13) Acer negundo Sarg. trees grown in a common garden in Salt Lake City, UT. The trees were established from cuttings collected where the growing season temperature averaged about 6.5 °C cooler than at the common garden. During May and June, stem sap flux (J_s) was similar between genders, but averaged 25% higher in males during the warmer months of July and August. Daytime canopy stomatal conductance (g_s) per unit leaf area was 12% higher in females in May : June, but was 11% higher in males in July : August. We combined measurements of sap flux–scaled transpiration with measurements of tree allometry and δ¹³C of leaf soluble sugars to estimate whole‐tree carbon assimilation (A_tree) and water use efficiency (WUE) (A_tree : E_tree). A_tree was similar between genders until late August when A_tree was 32% higher in male trees. A_tree : E_tree was on average 7% higher in females than in males during the growing season. Patterns of J_s, g_s, A_tree and A_tree : E_tree in the present study were in contrast to those previously reported for A. negundo genders under native growing season temperatures. Results suggest that the spatial segregation of the sexes could shift under global warming such that female plants lose their dominance in high‐resource habitats, and males increase their dominance in relatively lower‐resource habitats.     

广州地区荷木夜间树干液流补水的影响因子及其对蒸腾的贡献

明确树木夜间水分补充现象有助于提高总蒸腾量和冠层气孔导度估算的精确度,进一步认识冠层蒸腾与树干液流之间存在的时滞关系.本研究采用热消散探针法测定了广州地区的荷木树干液流密度,同步监测了主要的环境因子,从不同时间尺度分析了树干夜间液流的水分补充现象.结果表明:与白天相比,荷木夜间液流密度较小,旱季变化幅度比湿季大;夜间水分补充的时间段主要在前半夜(18:00-22:00);年内各季节夜间水分补充量之间没有显著差异,与环境因子之间的偏相关关系不显著,但与胸径、树高、冠幅、树干生物量、冠层生物量的回归曲线拟合很好,表明树形特征和生物量能更好地解释夜间补水的变化;各季节夜间水分补充量对总蒸腾量的贡献有显著差异,旱季明显高于湿季.     

The effect of tree height on crown level stomatal conductance

Variation in stomatal conductance is typically explained in relation to environmental conditions. However, tree height may also contribute to the variability in mean stomatal conductance. Mean canopy stomatal conductance of individual tree crowns (G_Si) was estimated using sap flux measurements in Fagus sylvatica L., and the hypothesis that G_Si decreases with tree height was tested. Over 13 d of the growing season during which soil moisture was not limiting, G_Si decreased linearly with the natural logarithm of vapour pressure deficit (D), and increased exponentially to saturation with photosynthetic photon flux density (Q_o). Under conditions of D = 1 kPa and saturating Q_o, G_Si decreased by approximately 60% with 30 m increase in tree height. Over the same range in height, sapwood‐to‐leaf area ratio (A_S:A_L) doubled. A simple hydraulic model explained the variation in G_Si based on an inverse relationship with height, and a linear relationship with A_S:A_L. Thus, in F. sylvatica, adjustments in A_S:A_L partially compensate for the negative effect of increased flow‐path length on leaf conductance. Furthermore, because stomata with low conductance are less sensitive to D, gas exchange of tall trees is reduced less by high D. Despite these compensations, decreasing hydraulic conductance with tree height in F. sylvatica reduces carbon uptake through a corresponding decrease in stomatal conductance.     

Nocturnal sap flow in the C<Subscript>3</Subscript>-CAM species, <Emphasis Type="Italic">Clusia minor</Emphasis>

Clusia minor L. is a C₃-CAM species in which Crassulacean acid Metabolism (CAM) is induced, among other factors, by water deficit. We propose that CAM induction by natural drought in C. minor shifts the sap flow pattern from daytime to a night-time one, and that the decreased osmotic potential due to increased malate content in droughted plants aids in the increase in nocturnal sap flow. In order to test these hypotheses, we followed for 2 years the seasonal changes in parameters of water relationships and sap flow velocity in one single, freestanding tree growing in Caracas. Leaf water and osmotic potential were measured psychrometrically, nocturnal proton accumulation by titration of aqueous leaf extracts and sap flow density with thermal dissipation probes. Leaf water, osmotic and turgor potential remained relatively high throughout the seasons. Nocturnal proton accumulation was nil under extreme drought or after frequent and heavy rains, and high after moderate rainfall. Estimated malate and citrate concentrations contributed up to 80 and 60%, respectively, of the value of osmotic potential. The shape of the daily courses of sap flow velocity varied seasonally, from mostly diurnal during the dry season to mostly nocturnal after a short dry spell during the rainy season, when nocturnal acid accumulation attained high values. There was a strong positive relationship between the proportion of the integrated sap flow courses corresponding to the night and dawn [H⁺] (r ² = 0.88). Increased nocturnal sap flow in the CAM stage of the tree of C. minor may be explained by a lower osmotic potential due to an increased acid concentration, together with increased stomatal aperture, as suggested by increased nocturnal acid accumulation probably due to nocturnal CO₂ fixation.     

Photosynthetic characteristics in canopies of Quercus rubra, Quercus prinus and Acer rubrum differ in response to soil water availability

Photosynthesis and related leaf characteristics were measured in canopies of co-occurring Quercus rubra L. (red oak), Quercus prinus L. (chestnut oak) and Acer rubrum L. (red maple) trees. Mature (20+ m tall) trees were investigated at sites of differing soil water availability within a catchment (a drier upper site and a wetter lower site). Leaf photosynthetic characteristics differed significantly between species and in response to site and position in the canopy. Photosynthetic capacity (A_max) was significantly greater at the wetter site in all canopy strata in A. rubrum but not in Q. rubra or Q. prinus. Our findings for A. rubrum are generally consistent with those predicting that species with higher specific leaf area (SLA) will have higher A_max per unit leaf nitrogen (N) and that species with leaves with lower SLA (e.g. Q. rubra and Q. prinus) will have shallower slopes of the A_max-N relationship. Importantly, the relationships between A_max and N_area (and by implication photosynthetic nitrogen-use efficiency, PNUE) differed in A. rubrum between the sites, with PNUE significantly lower at the drier site. The lower photosynthetic capacity and PNUE must substantially reduce carbon acquisition capacity in A. rubrum under these field conditions. Maximum stomatal conductance (g_smax) differed significantly between species, with g_smax greatest in Q. rubra and Q. prinus. In Q. rubra and Q. prinus, g_smax was significantly lower at the upper site than the lower site. There was no significant response of g_smax to site in A. rubrum. These stomatal responses were consistent with the C_i/C_a ratio, which was significantly lower in leaves of Q. rubra and Q. prinus at the upper site, but did not differ between sites in A. rubrum. Leaf '¹³C was significantly lower in A. rubrum than in either Q. rubra or Q. prinus at both sites. These findings indicate differences in stomatal behaviour in A. rubrum which are likely to contribute to lower water use efficiency at both sites. Our results support the hypothesis that the two Quercus species, in contrast to A. rubrum, maintain photosynthetic capacity at the drier site whilst minimising transpirational water loss. They also suggest, based primarily on physiological evidence, that the ability of A. rubrum to compete with other species of these deciduous forests may be limited, particularly in sites of low moisture availability and during low rainfall years.     

Changes with seasonal flooding in sap flow of the tropical flood-tolerant tree species, <Emphasis Type="Italic">Campsiandra laurifolia</Emphasis>

In order to determine how flooding affects sap flow and hydraulic conductivity of the tolerant species, Campsiandra laurifolia, trees growing in a tropical seasonally flooded forest in Venezuela were studied. We hypothesized that trees respond to rising-waters with a decrease in root-water absorption, caused by hypoxia, and stomatal conductance, and that this is reverted later on through a process of acclimation that involves improvement in water absorption. We followed the seasonal changes, of trees with the whole or part of the canopy exposed to air, in sap flow density, leaf stomatal conductance, leaf transpiration rate and xylem water potential. The highest daytime sap flow density occurred at noon and its proportion relative to the yearly maximum (drainage at falling-waters) was 41 (dry season), 15 (flooding by rising-waters for 2 weeks), 54 (2 months of flooding) and 41% (6 months of flooding). Since at rising-waters dawn xylem water potential remained high, it became apparent that the initial stages of flooding imposed a restriction to sap flow unrelated to water deficit. The decrease at rising-waters in highest daytime sap flow density was due to reduced leaf-specific hydraulic conductivity, whereas the recovery observed 1.5 months later was correlated to an increase in hydraulic conductivity, and attributed to acclimation. Sap flow density was highly and positively correlated with radiation at all seasons but rising-waters; also, the relationship with air water vapor saturation deficit was high and significant on dates other than at rising-waters. Results suggest that early flooding inhibited water absorption by roots and that this inhibition was overcome later on at a higher water column through an acclimation process involving the improvement of internal aeration by adventitious roots.     

Field water relations of a wet-tropical forest tree species,Pentaclethra macroloba (Mimosaceae)

Summary The water relations of Pentaclethra macroloba (Willd.) Kuntze, a dominant, shade-tolerant, tree species in the Atlantic lowlands of Costa Rica, were examined within the forest canopy. Pressure-volume curves and diurnal courses of stomatal conductance and leaf water potential were measured in order to assess differences in water relations between understory, mid-canopy and canopy leaves. Leaves in the canopy had the smallest pinnules but the largest stomatal frequencies and stomatal conductances of the three forest levels. Osmotic potentials at full turgidity decreased with height in the forest; in the canopy and midcanopy they were reduced relative to those in the understory just enough to balance the gravitational component of water potential. Consequently, maximum turgor pressures were similar for leaves from all three canopy levels. Bulk tissue elastic modulus increased with height in the canopy. Leaf water potentials were lowest in the canopy and highest in the understory, even when the gravitational component was added to mid-canopy and canopy values. As a result, minimum turgor pressures were also lowest in the canopy compared to those at lesser heights, and approached zero in full sunlight on clear days.Osmotic potentials at each canopy level were similar for both wet and dry season samples dates suggesting that seasonal osmotic adjustment does not occur. Despite lowered predawn water potentials during the dry season, turgor was maintained in the understory by reduced stomatal conductances.     

Photosynthesis and related physiological processes in two mangrove species,Rhizophora mucronata and Ceriops tagal,at Gazi Bay,Kenya

Measurements were carried out of the gas exchange properties (namely, photosynthesis, stomatal conductance and transpiration rates), water use efficiency and water relations of two mangrove species, Rhizophora mucronata and Ceriops tagal at Gazi Bay, Kenya. Rhizophora mucronata had significantly higher photosynthetic rates than C. tagal. Internal CO₂ concentrations were higher during the wet season than the dry season in both species. Gas exchange properties were correlated positively with photon flux density in both species. Leaf water potentials were highest during the morning and lowest at midday and were also highest in the lower canopy leaves in both species. The two mangrove species had conservative water use. Management potential for the East African mangroves based on the results of this study is suggested.