Hydrogen and oxygen isotope ratios of tree-ring cellulose for riparian trees grown long-term under hydroponically controlled environments

Saplings of three riparian tree species (alder, birch and cottonwood) were grown for over 5 months in a hydroponics system that maintained the isotopic composition of source water in six treatments, ranging from –120 to +180‰δD and –15 to +10‰δ¹⁸O. The trees were grown in two greenhouses maintained at 25°C and at either 40 or 75% relative humidity, creating differences in transpiration rates and leaf water isotopic evaporative enrichment. The cellulose produced in the annual growth ring was linearly related to source water with differences in both slope and offset associated with greenhouse humidity. The slope of the isotopic composition of source water versus tree-ring cellulose was less than 1 for both δD and δ¹⁸O indicating incomplete isotopic exchange of carbohydrate substrate with xylem water during cellulose synthesis. Tests using the outer portion of the tree-ring and new roots were similar and showed that the tree-ring values were representative of the cellulose laid down under the imposed environmental conditions. The fraction of H and O in carbohydrate substrate that isotopically exchange with medium water was calculated to be 0.36 and 0.42 respectively, and biochemical mechanisms for these observed fractions are discussed. A mechanistic model of the biochemical fractionation events for both δD and δ¹⁸O leading to cellulose synthesis was robust over the wide range of cellulose stable isotope ratios. The experimental results indicate that both water source and humidity information are indeed recorded in tree-ring cellulose. These results help to resolve some of the disparate observations regarding the interpretation of stable isotope ratios in tree-rings found in the literature. Received: 4 January 1999 / Accepted: 12 August 1999     

Leaf carbon isotope discrimination and nitrogen content for riparian trees along elevational transects

 Leaf carbon isotope discrimination (Δ), seasonal estimates of the leaf-to-air water vapor gradient on a molar basis (ω), and leaf nitrogen contents were examined in three riparian tree species (Populus fremontii, P. angustifolia, and Salix exigua) along elevational transects in northern and southern Utah USA (1500–2670 m and 600–1820 m elevational gradients, respectively). The ω values decreased with elevation for all species along transects. Plants growing at higher elevations exhibited lower Δ values than plants at lower elevations (P. fremontii, 22.9‰ and 19.5‰, respectively; P. angustifolia, 23.2‰ and 19.2‰, respectively; and S.␣exigua, 21.1‰ and 19.1‰, respectively). Leaf nitrogen content increased with elevation for all species, suggesting that photosynthetic capacity at a given intercellular carbon dioxide concentration was greater at higher elevations. Leaf Δ and nitrogen content values were highly correlated, implying that leaves with higher photosynthetic capacities also had lower intercellular carbon dioxide concentrations. No significant interannual differences were detected in carbon isotope discrimination. Received: 25 February 1996 / Accepted: 8 September 1996     

Seasonality in nutrient concentrations and stable isotope ratios of Halophila ovalis growing on the intertidal flat of SW Thailand

Halophila ovalis on the intertidal flat of SW Thailand maintains its presence under heavy grazing by dugongs; nevertheless, the site provides lower availability of nutrients in the water column to the seagrasses due to emersion during ebb tide. To examine the seasonality of the nutrient status of fast-growing H. ovalis in the intertidal flat, and to assess whether emersion at low tide affects the availability of nutrients and inorganic carbon, monthly changes in carbon (C), nitrogen (N), and phosphorus (P) concentrations of H. ovalis were determined. Carbon and nitrogen stable isotope ratios were also determined to examine the carbon and nitrogen sources. CNP concentrations of the leaves and petioles of H. ovalis at the intertidal flat were considerably lower than those of the same species previously reported from a subtidal environment throughout the observation period of 8 months. The δ¹³C of the above-ground parts of H. ovalis was lower than in previous reports, suggesting that CO₂ was sufficient for its growth. The CNP concentrations of H. ovalis on the intertidal flat increased after a decrease in grazing pressure by dugongs. It is suggested that H. ovalis sustained rapid growth on the intertidal flat by lowering its nutrient concentrations under heavy grazing by dugongs to maintain its presence. Received: March 2, 2001 / Accepted: June 6, 2001     

Disentangling effects of growth and nutritional status on seabird stable isotope ratios

A growing number of studies suggest that an individual’s physiology affects its carbon and nitrogen stable isotope signatures, obscuring a signal often assumed to be only a reflection of diet and foraging location. We examined effects of growth and moderate food restriction on red blood cell (RBC) and feather δ¹⁵N and δ¹³C in rhinoceros auklet chicks (Cerorhinca monocerata), a piscivorous seabird. Chicks were reared in captivity and fed either control (75 g/day; n = 7) or ~40% restricted (40 g/day; n = 6) amounts of high quality forage fish. We quantified effects of growth on isotopic fractionation by comparing δ¹⁵N and δ¹³C in control chicks to those of captive, non-growing subadult auklets (n = 11) fed the same diet. To estimate natural levels of isotopic variation, we also collected blood from a random sample of free-living rhinoceros auklet adults and chicks in the Gulf of Alaska (n = 15 for each), as well as adult feather samples (n = 13). In the captive experiment, moderate food restriction caused significant depletion in δ¹⁵N of both RBCs and feathers in treatment chicks compared to control chicks. Growth also induced depletion in RBC δ¹⁵N, with chicks exhibiting lower δ¹⁵N when they were growing the fastest. As growth slowed, δ¹⁵N increased, resulting in an overall pattern of enrichment over the course of the nestling period. Combined effects of growth and restriction depleted δ¹⁵N in chick RBCs by 0.92‰. We propose that increased nitrogen-use efficiency is responsible for ¹⁵N depletion in both growing and food-restricted chicks. δ¹⁵N values in RBCs of free-ranging auklets fell within a range of only 1.03‰, while feather δ¹⁵N varied widely. Together, our captive and field results suggest that both growth and moderate food restriction can affect stable isotope ratios in an ecologically meaningful way in RBCs although not feathers due to greater natural variability in this tissue.     

Branch sacrifice: cavitation-associated drought adaptation of riparian cottonwoods

In their native riparian zones (floodplains), Populus deltoides (prairie cottonwood) and P. fremontii (Fremont cottonwood) commonly experience substantial branch die-back. These trees occur in semi-arid areas of North America and unexpectedly given the dry regions, they are exceptionally vulnerable to xylem cavitation, drought-induced air embolism of xylem vessels. We propose that the vulnerability to cavitation and branch die-back are physiologically linked; drought-induced cavitation underlies branch die-back that reduces transpirational demand enabling the remaining shoot to maintain a favorable water balance. This proposal follows field observation along various western North American rivers as precocious branch senescence, the yellowing and death of leaves on particular branches during mid- to late summer, was common for P. deltoides and P. fremontii during hot and dry periods of low stream-flow. Branches displaying precocious senescence were subsequently dead the following year. The proposed association between cavitation, precocious senescence and branch die-back is also supported by experiments involving external pressurization of branches to about 2.5 MPa with a branch collar or through an adjacent cut-branch. The treatments induced xylem cavitation and increased leaf diffusive resistance (stomatal closure) that was followed by leaf senescence and branch death of P. deltoides. P. trichocarpa (black cottonwood) appeared to be less affected by the pressurization treatment and this species as well P. angustifolia (narrowleaf cottonwood) and P. balsamifera (balsam poplar) seldom display the patchy summer branch senescence typical of P. deltoides and P. fremontii. ’Branch sacrifice’ describes this cavitation-associated senescence and branch die-back that may provide a drought adaptation for the prairie and Fremont cottonwoods. Received: 13 May 1999 / Accepted: 4 November 1999     

Transpiration and whole-tree conductance in ponderosa pine trees of different heights

Changes in leaf physiology with tree age and size could alter forest growth, water yield, and carbon fluxes. We measured tree water flux (Q) for 14 ponderosa pine trees in two size classes (12 m tall and ∼40 years old, and 36 m tall and ∼ 290 years old) to determine if transpiration (E) and whole-tree conductance (g _t) differed between the two sizes of trees. For both size classes, E was approximately equal to Q measured 2 m above the ground: Q was most highly correlated with current, not lagged, water vapor pressure deficit, and night Q was <12% of total daily flux. E for days 165–195 and 240–260 averaged 0.97 mmol m^–2 (leaf area, projected) s^–1 for the 12-m trees and 0.57 mmol m^–2 (leaf area) s^–1 for the 36-m trees. When photosynthetically active radiation (I _P) exceeded the light saturation for photosynthesis in ponderosa pine (900 μmol m^–2 (ground) s^–1), differences in E were more pronounced: 2.4 mmol m^–2 (leaf area) s^–1 for the 12-m trees and 1.2 mmol m^–2 s^–1 for the 36-m trees, yielding g _t of 140 mmol m^–2 (leaf area) s^–1 for the 12-m trees and 72 mmol m^–2 s^–1 for the 36-m trees. Extrapolated to forests with leaf area index =1, the 36-m trees would transpire 117 mm between 1 June and 31 August compared to 170 mm for the 12-m trees, a difference of 15% of average annual precipitation. Lower g _t in the taller trees also likely lowers photosynthesis during the growing season. Received: 19 April 1999 / Accepted: 23 March 2000     

Scaling of tree height and trunk diameter as a function of ring number

 Phipps (1967) showed, in suppressed maples and oaks, that on average the cross-sectional area of successive tree rings is independent of ring number. From that empirical result I argued that mean ring width and mean ring diameter scale as the inverse square root and square root, respectively, of ring number (Prothero 1997). Here I give an illustrative argument as to how macroscopic trunk variables may be related quantitatively to microscopic ring variables. First, evidence is presented consistent with the theory of Greenhill (1881) and the empirical evidence of McMahon (1973) that tree height in a diversity of species scales as about the two-thirds power of trunk diameter. Here I report further evidence bearing on the same question. From these combined results I suggest that tree height, and by implication trunk height, and other parameters governing macroscopic tree trunk proportions, can all be expressed as powers of a microscopic variable, namely ring number, where the scaling exponents are all proper fractions. I suggest that at least one of these relationships is, in principle, non-adaptive. Received: 9 June 1998 / Accepted: 4 February 1999     

Hydrogen exchange during cellulose synthesis distinguishes climatic and biochemical isotope fractionations in tree rings

The abundance of the hydrogen isotope deuterium (D) in tree rings is an attractive record of climate; however, use of this record has proved difficult so far, presumably because climatic and physiological influences on D abundance are difficult to distinguish. Using D labelling, we created a D gradient in trees. Leaf soluble sugars of relatively low D abundance entered cellulose synthesis in stems containing strongly D-labelled water. We used nuclear magnetic resonance (NMR) spectroscopy to quantify D in the C-H groups of leaf glucose and of tree-ring cellulose. Ratios of D abundances of individual C-H groups of leaf glucose depended only weakly on leaf D labelling, indicating that the D abundance pattern was determined by physiological influences. The D abundance pattern of tree-ring cellulose revealed C-H groups that exchanged strongly (C(2)-H) or weakly (C(6)-H2) with water during cellulose synthesis. We propose that strongly exchanging C-H groups of tree-ring cellulose adopt a climate signal stemming from the D abundance of source water. C-H groups that exchange weakly retain their D abundance established in leaf glucose, which reflects physiological influences. Combining both types of groups may allow simultaneous reconstruction of climate and physiology from tree rings.     

Establishing pathways of energy flow for insect predators using stable isotope ratios: field and laboratory evidence

 Quantifying pathways of energy transfer between plants, pests, and beneficial insects is a necessary step toward maintaining pest stable agroecosystems in the absence of chemical subsidies. A diet switching experiment utilizing a predatory ladybird beetle, Hippodamia variegata (Goeze), evaluated the use of naturally occurring stable C and N isotopes as an economically feasible and safe method for quantifying pathways of energy flow within agroecosystems. Stable isotope values of the ladybird beetle Coleomegilla maculata lengi (Timberlake) collected from an agroecosystem were used to estimate the relative amount of C and N derived from agricultural plants and incorporated by ladybird beetles based on mass balance equations. At the beginning of the diet-switching experiment δ¹³C and δ¹⁵N values of H. variegata (–12.0‰ and 6.3‰, respectively) differed by –0.2‰ and 2.9‰ from the aphids that were provided exclusively as their diet. These data are consistent with previous estimates of trophic level isotope effects. After switching the diet of H. variegata to an alternative food, isotope values of H. variegata gradually shifted toward expected values for individuals fed this diet (–22.9‰ and 8.8‰ for δ¹³C and δ¹⁵N values, respectively). Isotope values of another ladybird beetle, C. maculata, collected from the field indicated that in May, alfalfa and maize (pollen) obtained in the previous year contributed 32% and 68% of the C or N to the diets of these individuals and in August, 52%, 6%, and 42% of the C or N assimilated by these insects was derived from alfalfa, wheat, and maize, respectively. These data are consistent with expectations based on the relative abundance of C. maculata in various crops during the season. The field and laboratory data are a clear indication that isotope values are sensitive to dietary changes on a relatively short time scale (days) and provide a strong basis for the use stable C and N isotope to trace energy flow patterns of these beneficial organisms within agroecosystems. Received: 17 November 1995 / Accepted: 20 June 1996     

Stable isotope analyses of otoliths in identification of hatchery origin of Atlantic salmon (Salmo salar) in Maine

We conducted stable oxygen and carbon isotope analyses for otoliths of Atlantic salmon (Salmo salar), in an attempt to develop a reference database on isotopic variability among private and federal hatcheries in Maine which currently support the salmon aquaculture industry and recovery of endangered populations. During the first phase of our study, we collected 40–50 sagittal otoliths of juvenile Atlantic salmon from each of the five hatcheries and analyzed for stable oxygen and carbon isotope ratios (¹⁸O/¹⁶O or δ¹⁸O, and ¹³C/¹²C or δ¹³C). Combination of δ¹⁸O and δ¹³C signatures in otoliths showed that the five hatcheries can be clearly separated and chemically distinguished. By identifying stable isotopic variations of otoliths from different hatchery settings, we were able to establish some isotopic criteria or standards to assign a likelihood that an individual Atlantic salmon came from a specific hatchery within the reference database. If successful, a diagnostic tool that can provide definitive information on identification of the hatchery origin could serve as a novel marking technique, and the chemical method may provide a more effective alternative to DNA analysis for mixed stocks. Overall our isotopic data from otoliths support the hypothesis that there are detectable differences between the five hatcheries, and multiple statistical analyses indicated that we can correctly distinguish individual Atlantic salmon into a hatchery with high confidence.     

Trends in plant carbon concentration and plant demand for N throughout this century

 Atmospheric CO₂ concentration has increased by 25% over the preindustrial level. A parallel increase in C concentration and decreases in N concentration and δ¹³C of plants grown throughout this century have been observed in plant specimens stored in herbaria. We tested our previous results in a study of 12 more species collected in the western Mediterranean throughout this century (1920–1930, 1945–1955, and 1985–1990) and tree rings of Quercus pubescens from the same area. These changes were accompanied by apparent increases in condensed tannin concentration. A decreasing trend in δ¹⁵N both in herbarium material and tree rings was also found, indicating that ecosystems might cope with higher plant N demand by decreasing N losses and increasing N fixation and mineralization. These results may contribute to a better understanding of the effects of global change on carbon and nitrogen cycling. Received: 12 November 1995 / Accepted: 17 May 1996     

Hydrogen, oxygen, and carbon isotope ratios of cellulose from submerged aquatic crassulacean Acid metabolism and non-crassulacean Acid metabolism plants

Isotope ratios of cellulose and cellulose nitrate from aquatic Crassulacean acid metabolism (CAM) and non-CAM plants were determined. Cellulose oxygen istope ratios for all plants that grew together were virtually identical, whereas large differences were observed for hydrogen isotope ratios of cellulose nitrate between CAM and non-CAM plants. Carbon isotope ratios of cellulose nitrate did not differentiate CAM from non-CAM plants.     

Effects of nutrient enrichment on C and N stable isotope ratios of invertebrates,fish and their food resources in boreal streams

Carbon and nitrogen stable isotopes are frequently used to study energy sources and food web structure in ecosystems, and more recently, to study the effects of anthropogenic stress on aquatic ecosystems. We investigated the effect of nutrient enrichment on δ¹³C and δ¹⁵N in fine (FPOM), coarse (CPOM) particulate organic matter, periphyton, invertebrates and fish in nine boreal streams in south-central Sweden. In addition, we analysed the diet of benthic consumers using stable isotope data. Increases in δ¹⁵N of periphyton (R ² = 0.88), CPOM (0.78), invertebrates (0.92) and fish (0.89) were related to nutrient enrichment. In contrast, δ¹³C signatures did not change along the nutrient gradient. Our results show that δ¹⁵N has potential as a sensitive indicator of nutrient enrichment in boreal streams. Carbon and nitrogen isotopes failed to elucidate putative diets of selected aquatic consumers. Indeed, comparison of low- and high-impact sites showed that δ¹³C of many consumers were found outside the ranges of basal resource δ¹³C. Moreover, ranges of basal resource δ¹³C and δ¹⁵N overlapped at both low and high sites, making discrimination between the importance of allochthonous and autochthonous production difficult. Our findings show that a fractionation rate of 3.4‰ is not always be appropriate to assess trophic interactions, suggesting that more studies are needed on fractionation rates along gradients of impairment. Handling editor: M. Power     

Resonance Raman as a direct probe for the catalytic mechanism of molybdenum oxotransferases

 Recent studies of human sulfite oxidase and Rhodobacter sphaeroides DMSO reductase have demonstrated the ability of resonance Raman to probe in detail the coordination environment of the Mo active sites in oxotransferases via Mo=O, Mo-S(dithiolene), Mo-S(Cys) or Mo-O(Ser), dithiolene chelate ring and bound substrate vibrations. Furthermore, the ability to monitor the catalytically exchangeable oxo group via isotopic labeling affords direct mechanistic information and structures for the catalytically competent Mo(IV) and Mo(VI) species. The results clearly demonstrate that sulfite oxidase cycles between cis–di-oxo-Mo(VI) and mono-oxo-Mo(IV) states during catalytic turnover, whereas DMSO reductase cycles between mono-oxo-Mo(VI) and des-oxo-Mo(IV) states. In the case of DMSO reductase, ¹⁸O-labeling experiments have provided the first direct evidence for an oxygen atom transfer mechanism involving an Mo=O species. Of particular importance is that the active-site structures and detailed mechanism of DMSO reductase in solution, as determined by resonance Raman spectroscopy, are quite different to those reported or deduced in the three X-ray crystallographic studies of DMSO reductases from Rhodobacter species. Received: 16 June 1997 / Accepted: 20 August 1997     

Effect of host shoot clipping on carbon and nitrogen sources for arbuscular mycorrhizal fungi

The effect of clipping of the host-plant shoot on the sources of carbon and nitrogen for the arbuscular mycorrhizal (AM) fungus Gigaspora margarita was determined by measuring ¹³C in spores and hyphae in cocultures of C₃ and C₄ plants and by differential ¹⁵N labeling. C₃ and C₄ plants, which have different δ¹³C values, were grown in the same container separated by a series of hyphal compartments. The C₃ and C₄ plants were applied with ¹⁴N- and ¹⁵N-urea, respectively. After clipping of the C₃ shoots, spore δ ¹³C gradually approached that of the C₄ roots. Hyphal δ ¹³C paralleled that of spores. Spore % ¹⁵N was similar to that of mineral N in the C₄ plant compartment. Thus C in G. margarita coming from the clipped plants decreased with time. This demonstrates that C in AM fungi comes from living plants, whilst the N in spores comes mostly from the soil. Accepted: 28 November 2000     

Influence of leaf size on tree architecture: first branch height and crown dimensions in tropical rain forest trees

 First branch height is an important attribute of sapling architecture, as it defines the height at which prolonged lateral growth is possible. First branch height, measured on saplings of 70 species in tropical rain forests of Australia, Costa Rica, Panama, and Sabah, Malaysia, was highly correlated with leaf blade and petiole length. The observed relationship, first branch height ∝ blade length × (petiole length)^0.5, implies that the ratio of first branch height to blade length increases somewhat with increasing leaf size, among species with a given ratio of petiole to blade length. Orthotropic species, with more or less radially symmetric arrangements of leaves on ascending axes, had a mean first branch height of 7x that observed for plagiotropic species, with planar leaf arrangements. The greater first branch height of orthotropic species was associated with their larger leaves and longer petioles. Plagiotropic species had wider crowns than orthotropic species in the sapling stage, as assessed at the Costa Rican site. Thus, leaf dimensions influence the dynamics of crown construction (or visa versa), as well as affecting leaf energy balance and gas exchange. Received: 5 September 1997 / Accepted: 3 March 1998     

Foliar carbon isotope discrimination in Larix species and sympatric evergreen conifers: a global comparison

Larches (Larix spp.), deciduous conifers, occur in the northern hemisphere in cold-temperate and boreal climates – an environment normally thought to favor evergreen tree species. We compare foliar carbon isotope discrimination (Δ), instantaneous water use efficiency, total foliar nitrogen concentration, and specific leaf area (for a subset of sites) between Larix spp. and co-occurring evergreen conifers at 20 sites throughout the natural range of larches. Except for Larix occidentalis in the xeric Intermountain West, USA, Δ is significantly (P < 0.05) greater for larches than co-occurring evergreen conifers at 77% of the sites, suggesting that larches use water less efficiently. At elevations greater than 3000 m, the Δ of Larix spp. and co-occurring conifers converge, suggesting that water is not the limiting resource. Foliar nitrogen concentration and specific leaf area are two ecophysiological characteristics that are positively correlated with high photosynthetic capacity. Foliar nitrogen concentration is significantly greater for larches than evergreen conifers at 88% of the sites and specific leaf area is approximately three times greater for larches than co-occurring conifers. Future studies should examine the potential effect that global warming may have on the distribution of larch forests because the water use efficiency of larches is commonly less than co-occurring evergreen conifers and the boreal and high-latitude environments are likely to experience the greatest climate warming. Received: 23 May 1997 / Accepted: 28 October 1997     

Relative humidity- and ABA-induced variation in carbon and oxygen isotope ratios of cotton leaves

Cotton (Gossypium hirsutum L. cv. CS50) plants were grown at two levels of relative humidity (RH) and sprayed daily with abscisic acid (ABA) at four concentrations. Plants grown at lower humidity had higher transpiration rates, lower leaf temperatures and lower stomatal conductance. Plant biomass was also reduced at low humidity. Within each humidity environment, increasing ABA concentration generally reduced stomatal conductance, evaporation rates, superficial leaf density and plant biomass, and increased leaf temperature and specific leaf area. As expected, decreased stomatal conductance resulted in decreased carbon isotope discrimination in leaf material ( Δ ¹³C_l). Plants grown at low humidity were more enriched in ¹⁸O than those grown at high RH, as theory predicts. Within each humidity environment, increasing ABA concentration increased oxygen isotope enrichment of leaf cellulose ( Δ ¹⁸O_c) and whole‐leaf tissue ( Δ ¹⁸O_l). Values of Δ ¹³C_l and Δ ¹⁸O_l predicted by theoretical models were close to those observed, accounting for 79% of the measured variation in Δ ¹³C_l and 95% of the measured variation in Δ ¹⁸O_l. Supporting theory, Δ ¹³C_l and Δ ¹⁸O_l in whole‐leaf tissue were negatively related.     

Changes in drought response strategies with ontogeny in Quercus rubra: implications for scaling from seedlings to mature trees

We investigated scaling of physiological parameters between age classes of Quercus rubra by combining in situ field measurements with an experimental approach. In the in situ field study, we investigated changes in drought response with age in seedlings, juveniles, and mature trees of Q. rubra. Throughout the particularly dry summer of 1995 and the unusually wet summer of 1996 in New England, we measured water potential of leaves (Ψ_Leaf) and gas exchange of plants at three sites at the Harvard Forest in Petersham, Massachusetts. In order to determine what fraction of the measured differences in gas exchange between seedlings and mature trees was due to environment versus ontogeny, an experiment was conducted in which seedlings were grown under light and soil moisture regimes simulating the environment of mature trees. The photosynthetic capacity of mature trees was three-fold greater than that of seedlings during the wet year, and six-fold greater during the drought year. The seedling experiment demonstrated that the difference in photosynthetic capacity between seedlings and mature trees is comprised equally of an environmental component (50%) and an ontogenetic component (50%) in the absence of water limitation. Photosynthesis was depressed more severely in seedlings than in mature trees in the drought year relative to the wet year, while juveniles showed an intermediate response. Throughout the drought, the predawn leaf water potential (Ψ_PD) of seedlings became increasingly negative (–0.4 to –1.6 MPa), while that of mature trees became only slightly more negative (–0.2 to –0.5 MPa). Again, juveniles showed an intermediate response (–0.25 to –0.8 MPa). During the wet summer of 1996, however, there was no difference in Ψ_PD between seedlings, juveniles and mature trees. During the dry summer of 1995, seedlings were more responsive to a major rain event than mature trees in terms of Ψ_Leaf , suggesting that the two age classes depend on different water sources. In all age classes, instantaneous measurements of intrinsic water use efficiency (WUE_i), defined as C assimilation rate divided by stomatal conductance, increased as the drought progressed, and all age classes had higher WUE_i during the drought year than in the wet year. Mature trees, however, showed a greater ability to increase their WUE_i in response to drought. Integrated measurements of WUE from C isotope discrimination (Δ) of leaves indicated higher WUE in mature trees than juveniles and seedlings. Differences between years, however, could not be distinguished, probably due to the strong bias in C isotope fractionation at the time of leaf production, which occurred prior to the onset of drought conditions in 1995. From this study, we arrive at two main conclusions: Received: 14 July 1999 / Accepted: 10 January 2000     

Photosynthesis rates of selected tree species in lowland dipterocarp rainforest of Sabah, Malaysia

 Diurnal courses of net photosynthesis, transpiration and water potential of leaves of ten woody species from the natural lowland dipterocarp forests in Sabah (North Borneo, Malaysia) and one exotic tree species were studied in the field. The indigenous species represent different ecological niches and successional stages in the various layers of the dipterocarp forest, such as pioneers, trees of the understorey or main canopy and emergents. Diurnal changes in CO₂ exchange and transpiration reflected primarily differences in irradiance. The diurnal courses of water potential mainly tracked the rate of transpiratory water loss. Light-dependency describes most of the diurnal variations of leaves’ gas exchange. Light response curves of net photosynthesis of the investigated species of the Dipterocapaceae were almost equal (light saturated assimilation rate, A_max: 5.0–7.2 μmol CO₂ m^–2 s^–1), while those of the other species exhibited remarkable differences (A_max: 5.5–14.2 μmol CO₂ m^–2 s^–1). Leaf area, chlorophyll content and specific leaf dry weight as the reference parameters for assimilation gave a general ranking of the A_max, which is highest for the pioneering species, less for the understorey trees and lowest for emergents. Light compensation points and light saturation of net photosynthesis were attained mainly between 6 and 9 μmol photons m^–2 s^–1 and between 230 and 534 μmol photons m^–2 s^–1, respectively, but were higher for pioneering species. Photosynthetic performance may be a diagnostic feature of the successional and ecological status of species, i.e. to characterize pioneering species from understorey species or from emergents of the dipterocarp forest. Received: 3 March 1997 / Accepted: 15 December 1997