Relationships of stable carbon isotopes, plant water potential and growth: an approach to asses water use efficiency and growth strategies of dry land agroforestry species

The relationships between annual wood stable carbon isotope composition (δ¹³C), dry season midday plant water potential, and annual growth rate were investigated to asses the ability of agroforestry species to adapt to climate changes. 6–8 stem disks from four co-occurring species (Acacia senegal, A. seyal, A. tortilis and Balanites aegyptiaca) were collected for radial growth measurements using tree-ring analysis spanning 1930–2003. Annual δ¹³C was measured on three tree disks per species for the period 1970–2002. Midday plant water potential was measured during the dry season. Annual radial growth and midday plant water potential ranged from 0.27 to 9.12 mm and −1.0 to −5.0 MPa, respectively, with statistically significant differences. After correcting annual wood δ¹³C for atmospheric changes in δ¹³C, carbon isotopic composition ranged from −22.22 to −26.58‰. Relationships between δ¹³C, radial growth and plant water potentials revealed the interaction of water availability, stomatal conductance, δ¹³C values and growth. Two contrasting water use strategies and competitive advantages can be distinguished. Species with lower mean δ¹³C values (A. senegal and A. seyal) show high plant water potential and, hence, better growth during moist years. Thus, they indicate low water use efficiency (WUE) and opportunistic water use strategy. On the other hand, species with lower water potentials (A. tortilis and B. aegyptiaca) showed relative better growth performance and less increase in δ¹³C in drought years, reflecting their high WUE and conservative water use strategy. These results suggest that δ¹³C in tree rings can be useful in estimating historic changes in plant WUE and hence in screening drought tolerant species in the face of expected climate changes, as well as for assessing the functional diversity and risk reduction in mixed vegetation.     

��13C signature of tree rings and radial increment of Fagus sylvatica trees as dependent on tree neighborhood and climate

We conducted dendroecological analyses in 80-year-long tree ring chronologies to detect neighborhood effects (competition intensity, species identity) on the δ¹³C signature of tree rings and radial stem increment of Fagus sylvatica trees growing either in monospecific or mixed patches of a temperate forest. We hypothesized that tree ring δ¹³C is a more sensitive indicator of neighborhood effects and the impact of climate variability on growth than is ring width. We found a closer correlation of summer precipitation to δ¹³C than to ring width. While the ring width showed a decline over the test period (1926–2005), the mean curve of δ¹³C increased until the mid of the 1970s, remained high until about 1990, and markedly decreased thereafter. Possible explanations related to ontogeny and environmental change (‘age effect’ due to canopy closure; elevated atmospheric SO₂ concentrations in the 1960s–1980s) are discussed. Beech target trees surrounded by many allospecific trees had a significantly lower mean δ¹³C in the period 1926–1975 than beech with predominantly or exclusively conspecific neighborhood, possibly indicating a more favorable water supply of beech in diverse stands. Contrary to expectation, trees subject to more intense competition by neighboring trees (measured by Hegyi’s competition index) had lower δ¹³C values in their tree rings, which is thought to reflect denser canopies being linked to increased shading. We conclude that tree ring δ¹³C time series represent combined archives of climate variability, stand history and neighborhood effects on tree physiology and growth that may add valuable information to that obtained from conventional tree ring analysis.     

Stable oxygen isotopes (δ 18O) in Austrocedrus chilensis tree rings reflect climate variability in northwestern Patagonia, Argentina

The stable oxygen isotope (δ ¹⁸O) composition of Austrocedrus chilensis (D. Don) Endl. (Cupressaceae) tree rings potentially provide retrospective views of changes in environment and climate in the semi-arid lands of Patagonia. We report the development of the first annually resolved δ ¹⁸O tree-ring chronology obtained from natural forests of the foothills of the northwestern Patagonian Andes. The isotope record spans between 1890 and 1994 AD. We explore the probable links between this record and the climate of the region. Air temperatures during summer conditions are significantly, but not strongly, inversely correlated with annual δ ¹⁸O values from Austrocedrus tree rings. The strongest correlations are between the southern oscillation index (SOI) and the tree rings. The existence of millennial-age Austrocedrus trees in northern Patagonia provides interesting possibilities for examining these climate-related isotopic signals over most of the last 1,000 years.     

Detecting invisible growth rings of trees in seasonally dry forests in Thailand: isotopic and wood anatomical approaches

We measured radial variation of carbon isotope composition and vessel traits in tree species in seasonally dry forests of Northeast Thailand to explore a more reliable and amenable method of tropical dendrochronology for trees that lack visually detectable and consistent growth rings. Six Dipterocarpaceae species (3 Shorea, 2 Dipterocarpus, and 1 Hopea species) with indistinct or irregular growth rings and teak (Tectona grandis), a species which forms distinct growth rings, were examined. The δ¹³C value variations in all species showed annual cyclicity. Dipterocarpaceae species usually marked the lowest values of δ¹³C in the middle of the growing season, whereas teak had the lowest values at nearly the end of the growing season. Since the growing season of the species examined almost corresponds to the rainy season in the study area, the δ¹³C variation was likely caused by the change in moisture availability. The different variation pattern of teak was attributable to its stronger dependence on ¹³C-enriched reserved material early in the growing season. Changes in tree vessel traits for all species examined also showed annual cyclicity. Dipterocarpaceae species showed significant correlation between δ¹³C values and vessel measurements. Vessel lumen (mean area, tangential and radial diameter, and proportion of total area) had a negative correlation, whereas vessel frequency showed a positive correlation. The correlations indicated that changes in vessel traits were caused by the seasonal variation of moisture available to the trees. Thus, we concluded that methods using wood anatomy, as well as δ¹³C, have great potential for use as tools in tropical dendrochronology within the context of seasonal climate.     

Vegetation control effects on untreated wood,crude cellulose and holocellulose δ<Superscript>13</Superscript>C of early and latewood in 3- to 5-year-old rings of Douglas-fir

The stable carbon (C) composition of tree rings expressed as δ¹³C, is a measure of intrinsic water-use efficiency and can indicate the occurrence of past water shortages for tree growth. We examined δ¹³C in 3- to 5-year-old rings of Douglas-fir (Pseudotsuga menziesii (Mirb) Franco) trees to elucidate if decreased water supply or uptake was a critical factor in the observed growth reduction of trees competing with understory herb and shrub vegetation compared to those growing without competition. We hypothesized that there would be no differences in δ¹³C of earlywood in trees growing in plots with competing vegetation and those in plots receiving complete vegetation control during 5 years because earlywood formed early in the growing season when soil water was ample. We also hypothesized that δ¹³C in latewood which was formed during the later half of the growing season when precipitation was low, would be greater (less negative) in trees in plots without vegetation control. We then separated early and latewood from rings for three consecutive years and analyzed their δ¹³C composition. No significant differences in earlywood δ¹³C in years 3–5 were observed for trees in the two vegetation control treatments. δ¹³C of untreated latewood separated from wood cores was greater in 4- and 5-year-old rings of trees growing with competing vegetation compared to trees growing without vegetation competition (i.e., −25.5 vs. −26.3‰ for year 4, and −26.1 vs. −26.8‰ for year 5). Results suggest that water shortages occurred in Douglas-fir trees on this coastal Washington site in the latewood-forming portion of the growing season of years 4 and 5 in the no-vegetation control treatment. We also compared δ¹³C from untreated wood, crude cellulose extracted with the Diglyme–HCl method, and holocellulose extracted with toluene–ethanol to see if the extraction method would increase the sensitivity of the analysis. δ¹³C values from the two extraction methods were highly correlated with those from untreated samples (r ² = 0.97, 0.98, respectively). Therefore, using untreated wood would be as effective as using crude cellulose or holocellulose to investigate δ¹³C patterns in young Douglas-fir.     

Hydrogen and oxygen isotope ratios of tree ring cellulose for field-grown riparian trees

The isotopic composition of tree ring cellulose was obtained over a 2-year period from small-diameter riparian-zone trees at field sites that differed in source water isotopic composition and humidity. The sites were located in Utah (cool and low humidity), Oregon (cool and high humidity), and Arizona (warm and low humidity) with source water isotope ratio values of –125/–15‰ (δD/δ¹⁸O), –48/–6‰, and –67/–7‰, respectively. Monthly environmental measurements included temperature and humidity along with measurements of the isotope ratios in atmospheric water vapor, stream, stem, and leaf water. Small riparian trees used only stream water (both δD and δ¹⁸O of stem and stream water did not differ), but δ values of both atmospheric water vapor and leaf water varied substantially between months. Differences in ambient temperature and humidity conditions between sites contributed to substantial differences in leaf water evaporative enrichment. These leaf water differences resulted in differences in the δD and δ¹⁸O values of tree ring cellulose, indicating that humidity information was recorded in the annual rings of trees. These environmental and isotopic measurements were used to test a mechanistic model of the factors contributing to δD and δ¹⁸O values in tree ring cellulose. The model was tested in two parts: (a) a leaf water model using environmental information to predict leaf water evaporative enrichment and (b) a model describing biochemical fractionation events and isotopic exchange with medium water. The models adequately accounted for field observations of both leaf water and tree ring cellulose, indicating that the model parameterization from controlled experiments was robust even under uncontrolled and variable field conditions. Received: 7 April 1999 / Accepted: 8 December 1999     

Temperature versus species-specific influences on the stable oxygen isotope ratio of tree rings

Stable isotopic ratios integrate ecosystem variability while reflecting change in both environmental and biological processes. At sites, where climate does not strongly limit tree growth, co-occurring trees may display large discrepancies in stable oxygen isotopic ratios (δ¹⁸O) due to the interplay between biological processes (competition for light and nutrients, individual tree physiology, etc.) and climate. For a better quantification of the isotope variability within and among trees, the climatic and/or individual tree effects on seasonal δ¹⁸O variations in precipitation, soil water, leaf water and leaf organic material (whole leaf, cellulose and starch) and annual δ¹⁸O variations in tree-ring cellulose for Fagus sylvatica (Fs), Quercus robur (Qr), Carpinus betulus (Cb) and Pinus sylvestris (Ps) were studied in a mature temperate forest in Switzerland, using a mixed linear regression model technique. Furthermore, the influence of environmental factors on δ¹⁸O was assessed by means of three common isotope fractionation models. Our statistical analysis showed that except for Ps, a greater portion of δ¹⁸O variance in leaf compounds can be explained by individual tree effects, compared to temperature. Concerning tree-ring cellulose, only Fs and Ps show a significant temperature signal (maximum 12% of the variance explained), while the individual tree effect significantly explains δ¹⁸O for all species for a period of 38 years. Large species differences resulted in a limited ability of the isotope fractionation models to predict measured values. Overall, we conclude that in a diverse mixed forest stand, individual tree responses reduce the potential extraction of a temperature signal from δ¹⁸O.     

Stable isotope composition of <Emphasis Type="Italic">Chara rudis</Emphasis> incrustation in Lake Jasne,Poland

Stable isotope composition (δ¹³C and δ¹⁸O) was analysed in mineral incrustation of Chara rudis and surrounding waters. This macroalga forms dense and extensive charophyte meadows and may significantly contribute to the calcium carbonate precipitation and deposition of marl lake sediments. The study aimed to find out if charophyte calcium carbonate was precipitated in an isotopic equilibrium with lake water and if the precipitation was related to the environmental conditions. Two apical internodes of 10 individuals of C. rudis were collected monthly between June and late October 2008 at three permanent study sites (1.0 m, 1.5 m and 2.0 m deep) in a small (15.1 ha) and shallow (mean depth: 4.3 m) mid-forest lake with extensively developed charophyte meadows (Lake Jasne, mid-Western Poland). Basic physical–chemical analyses were performed at each study site, and water samples for further laboratory determinations, including stable isotope analyses, were collected from the above searched C. rudis stands and, simultaneously, at three comparative sites in the macrophyte-free pelagial. The difference in δ¹³C between incrustation and water from above C. rudis exceeded 2‰ V-PDB at each site. In the case of δ¹⁸O, it exceeded 2‰ between July and September. Accordingly, it is postulated that calcium carbonate was not precipitated in an isotopic equilibrium with lake water. Incrustation was enriched in heavier carbon isotope, ¹³C, and water was enriched in ¹⁸O. δ¹³C of incrustation and DIC were positively correlated, whereas negative relation was found between δ¹⁸O of incrustation and water. Several dependencies were found with water chemistry above the plants. The content of mineral incrustation in Chara dry weight had negative influence on the δ¹⁸O but not on the δ¹³C. Community depth, structure and PVI had no effect. No significant differences appeared between isotope composition in the pelagic zone and Chara stands.     

Spatial patterns of foliar stable carbon isotope compositions of C<Subscript>3</Subscript> plant species in the Loess Plateau of China

The spatial pattern of foliar stable carbon isotope compositions (δ¹³C) of dominant species and their relationships with environmental factors in seven sites, Yangling, Yongshou, Tongchuan, Fuxian, Ansai, Mizhi and Shenmu, standing from south to north in the Loess Plateau of China, was studied. The results showed that in the 121 C₃ plant samples collected from the Loess Plateau, the foliar δ¹³C value ranged from −22.66‰ to −30.70‰, averaging −27.04‰. The foliar δ¹³C value varied significantly (P<0.01) among the seven sites, and the average δ¹³C value increased by about 1.69‰ from Yangling in the south to Shenmu in the north as climatic drought increased. There was a significant difference in foliar δ¹³C value among three life-forms categorized from all the plant samples in the Loess Plateau (P<0.001). The trees (−26.74‰) and shrubs (−26.68‰) had similar mean δ¹³C values, both significantly (P<0.05) higher than the mean δ¹³C value of herbages (−27.69‰). It was shown that the trees and shrubs had higher WUEs and employed more conservative water-use patterns to survive drier habitats in the Loess Plateau. Of all the C₃ species in the Loess Plateau, the foliar δ¹³C values were significantly and negatively correlated with the mean annual rainfall (P<0.001) and mean annual temperature (P<0.05), while being significantly and positively correlated with the latitude (P<0.001) and the annual solar radiation (P<0.01). In general, the foliar δ¹³C values increased as the latitude and solar radiation increased and the rainfall and temperature decreased. The annual rainfall as the main influencing factor could explain 13.3% of the spatial variations in foliar δ¹³C value. A 100 mm increment in annual rainfall would result in a decrease by 0.88‰ in foliar δ¹³C values.     

Seasonal courses of key parameters of nitrogen,carbon and water balance in European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) grown on four different study sites along a European North–South climate gradient during the 2003 drought

During the growing season of the exceptionally dry and warm year 2003, we assessed seasonal changes in nitrogen, carbon and water balance related parameters of mature naturally grown European beech (Fagus sylvatica L.) along a North–South transect in Europe that included a beech forest stand in central Germany, two in southern Germany and one in southern France. Indicators for N balance assessed at all four sites were foliar N contents and total soluble non-protein nitrogen compounds (TSNN) in xylem sap, leaves and phloem exudates; C and water balance related parameters determined were foliar C contents, δ¹³C and δ¹⁸O signatures. Tissue sampling was performed in May, July and September. The N related parameters displayed seasonal courses with highest concentrations during N remobilization in May. Decreased total foliar N contents as well as higher C/N ratios in the stands in central Germany and southern France compared to the other study sites point to an impaired N nutrition status due to lower soil N contents and precipitation perception. TSNN concentrations in leaves and phloem exudates of all study sites were in ranges previously reported, but xylem sap content of amino compounds in July was lower at all study sites when compared to literature data (c. 1 μmol N mL⁻¹). In September, TSNN concentrations increased again at the two study sites in southern Germany after a rain event, whereas they remained constant at sites in central Germany and southern France which hardly perceived precipitation during that time. Thus, TSNN concentrations in the xylem sap might be indicative for water balance related N supply in the beech trees. TSNN profiles at all study sites, however, did not indicate drought stress. Foliar δ¹³C, but not foliar C and δ¹⁸O followed a seasonal trend at all study sites with highest values in May. Differences in foliar δ¹³C and δ¹⁸O did not reflect climatic differences between the sites, and are attributed to differences in altitude, photosynthesis and δ¹⁸O signatures of the water sources. Except of low TSNN concentrations in the xylem sap, no physiological indications of drought stress were detected in the trees analysed. We suppose that the other parameters assessed might not have been sensitive to the drought events because of efficient regulation mechanisms that provide a suitable physiological setting even under conditions of prolonged water limitation. The uniform performance of the trees from southern France and central Germany under comparably dry climate conditions denotes that the metabolic plasticity of mature beech from the different sites studied might be similar.     

Effects of climate, tree age, dominance and growth on δ15N in young pinewoods

Needles, annual rings from basal stem discs and bark of three dominant and three suppressed Pinus pinaster from a 12-year-old pine stand (naturally regenerated after a wildfire) were analysed to study the effects of climate, tree age, dominance, and growth on tree δ¹⁵N. Foliar-N concentration in dominant pines (0.780–1.474% N) suggested that soil N availability was sufficient, a circumstance that allowed isotopic discrimination by plants and (greater) differences in δ¹⁵N among trees. The δ¹⁵N decreases in the order wood (−0.20 to +6.12‰), bark (−1.84 to +1.85‰) and needles (−2.13 to +0.77‰). In all trees, before dominance establishment (years 1–8), the N stored in each ring displayed a decreasing δ¹⁵N tendency as the tree grows, which is mainly due to a more “closed” N cycle or an increasing importance of N sources with lower δ¹⁵N. After dominance establishment (years 9–12), wood δ¹⁵N values were higher in suppressed than in dominant trees (2.62 and 1.46‰, respectively; P < 0.01) while the reverse was true for needles and bark; simultaneously, the absolute amount of N stored by suppressed pines in successive rings decreased, suggesting a lower soil N assimilation. These results could be explained by lignification acting as major N source for needles in suppressed pines because products released and reallocated during lignification are ¹⁵N-depleted compared with the source. According to principal component analysis, wood δ¹⁵N appears associated with wood N concentration and precipitation during the growing season, but clearly opposed to age, basal area increment and mean temperature in spring and summer.     

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.     

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     

Intra-annual variability of anatomical structure and δ<Superscript>13</Superscript>C values within tree rings of spruce and pine in alpine,temperate and boreal Europe

Tree-ring width, wood density, anatomical structure and ¹³C/¹²C ratios expressed as δ¹³C-values of whole wood of Picea abies were investigated for trees growing in closed canopy forest stands. Samples were collected from the alpine Renon site in North Italy, the lowland Hainich site in Central Germany and the boreal Flakaliden site in North Sweden. In addition, Pinus cembra was studied at the alpine site and Pinus sylvestris at the boreal site. The density profiles of tree rings were measured using the DENDRO-2003 densitometer, δ¹³C was measured using high-resolution laser-ablation-combustion-gas chromatography-infra-red mass spectrometry and anatomical characteristics of tree rings (tracheid diameter, cell-wall thickness, cell-wall area and cell-lumen area) were measured using an image analyzer. Based on long-term statistics, climatic variables, such as temperature, precipitation, solar radiation and vapor pressure deficit, explained <20% of the variation in tree-ring width and wood density over consecutive years, while 29–58% of the variation in tree-ring width were explained by autocorrelation between tree rings. An intensive study of tree rings between 1999 and 2003 revealed that tree ring width and δ¹³C-values of whole wood were significantly correlated with length of the growing season, net radiation and vapor pressure deficit. The δ¹³C-values were not correlated with precipitation or temperature. A highly significant correlation was also found between δ¹³C of the early wood of one year and the late wood of the previous year, indicating a carry-over effect of the growing conditions of the previous season on current wood production. This latter effect may explain the high autocorrelation of long-term tree-ring statistics. The pattern, however, was complex, showing stepwise decreases as well as stepwise increases in the δ¹³C between late wood and early wood. The results are interpreted in the context of the biochemistry of wood formation and its linkage to storage products. It is clear that the relations between δ¹³C and tree-ring width and climate are multi-factorial in seasonal climates.     

Inter-annual and seasonal variability of radial growth, wood density and carbon isotope ratios in tree rings of beech (Fagus sylvatica) growing in Germany and Italy

We investigated the variability of tree-ring width, wood density and ¹³C/¹²C in beech tree rings (Fagus sylvatica L.), and analyzed the influence of climatic variables and carbohydrate storage on these parameters. Wood cores were taken from dominant beech trees in three stands in Germany and Italy. We used densitometry to obtain density profiles of tree rings and laser-ablation-combustion-GC-IRMS to estimate carbon isotope composition (δ ¹³C) of wood. The sensitivity of ring width, wood density and δ ¹³C to climatic variables differed; with tree-ring width responding to environmental conditions (temperature or precipitation) during the first half of a growing season and maximum density correlated with temperatures in the second part of a growing season (July–September). δ ¹³C variations indicate re-allocation and storage processes and effects of drought during the main growing season. About 20% of inter-annual variation of tree-ring width was explained by the tree-ring width of the previous year. This was confirmed by δ ¹³C of wood which showed a contribution of stored carbohydrates to growth in spring and a storage effect that competes with growth in autumn. Only mid-season δ ¹³C of wood was related to concurrent assimilation and climate. The comparison of seasonal changes in tree-ring maximum wood density and isotope composition revealed that an increasing seasonal water deficit changes the relationship between density and ¹³C composition from a negative relation in years with optimal moisture to a positive relationship in years with strong water deficit. The climate signal, however, is over-ridden by effects of stand density and crown structure (e.g., by forest management). There was an unexpected high variability in mid season δ ¹³C values of wood between individual trees (−31 to −24‰) which was attributed to competition between dominant trees as indicated by crown area, and microclimatological variations within the canopy. Maximum wood density showed less variation (930–990 g cm⁻³). The relationship between seasonal changes in tree-ring structure and ¹³C composition can be used to study carbon storage and re-allocation, which is important for improving models of tree-ring growth and carbon isotope fractionation. About 20–30% of the tree-ring is affected by storage processes. The effects of storage on tree-ring width and the effects of forest structure put an additional uncertainty on using tree rings of broad leaved trees for climate reconstruction.     

Water source partitioning among trees growing on shallow karst soils in a seasonally dry tropical climate

The sources of water used by woody vegetation growing on karst soils in seasonally dry tropical regions are little known. In northern Yucatan (Mexico), trees withstand 4–6 months of annual drought in spite of the small water storage capacity of the shallow karst soil. We hypothesized that adult evergreen trees in Yucatan tap the aquifer for a reliable supply of water during the prolonged dry season. The naturally occurring concentration gradients in oxygen and hydrogen stable isotopes in soil, bedrock, groundwater and plant stem water were used to determine the sources of water used by native evergreen and drought-deciduous tree species. While the trees studied grew over a permanent water table (9–20 m depth), pit excavation showed that roots were largely restricted to the upper 2 m of the soil/bedrock profile. At the peak of the dry season, the δ¹⁸O signatures of potential water sources for the vegetation ranged from 4.1 ± 1.1‰ in topsoil to −4.3 ± 0.1‰ in groundwater. The δ¹⁸O values of tree stem water ranged from −2.8 ± 0.3‰ in Talisia olivaeformis to 0.8 ± 1‰ in Ficus cotinifolia, demonstrating vertical partitioning of soil/bedrock water among tree species. Stem water δ¹⁸O values were significantly different from that of groundwater for all the tree species investigated. Stem water samples plotted to the right of the meteoric water line, indicating utilization of water sources subject to evaporative isotopic enrichment. Foliar δ¹³C in adult trees varied widely among species, ranging from −25.3 ± 0.3‰ in Enterolobium cyclocarpum to −28.7 ± 0.4‰ in T. olivaeformis. Contrary to initial expectations, data indicate that native trees growing on shallow karst soils in northern Yucatan use little or no groundwater and depend mostly on water stored within the upper 2–3 m of the soil/bedrock profile. Water storage in subsurface soil-filled cavities and in the porous limestone bedrock is apparently sufficient to sustain adult evergreen trees throughout the pronounced dry season.     

Association between tree-ring and needle δ<Superscript>13</Superscript>C and leaf gas exchange in <Emphasis Type="Italic">Pinus halepensis</Emphasis> under semi-arid conditions

Associations between δ¹³C values and leaf gas exchanges and tree-ring or needle growth, used in ecophysiological compositions, can be complex depending on the relative timing of CO₂ uptake and subsequent redistribution and allocation of carbon to needle and stem components. For palaeoenvironmental and dendroecological studies it is often interpreted in terms of a simple model of δ¹³C fractionation in C₃ plants. However, in spite of potential complicating factors, few studies have actually examined these relationships in mature trees over inter- and intra-annual time-scales. Here, we present results from a 4 years study that investigated the links between variations in leaf gas-exchange properties, growth, and dated δ¹³C values along the needles and across tree rings of Aleppo pine trees growing in a semi-arid region under natural conditions or with supplemental summer irrigation. Sub-sections of tissue across annual rings and along needles, for which time of formation was resolved from growth rate analyses, showed rapid growth and δ¹³C responses to changing environmental conditions. Seasonal cycles of growth and δ¹³C (up to ~4‰) significantly correlated (P<0.01) with photosynthetically active radiation, vapour pressure deficit, air temperature, and soil water content. The irrigation significantly increased leaf net assimilation, stomatal conductance and needle and tree-ring growth rate, and markedly decreased needle and tree-ring δ¹³C values and its sensitivity to environmental parameters. The δ¹³C estimates derived from gas-exchange parameters, and weighted by assimilation, compared closely with seasonal and inter-annual δ¹³C values of needle- and tree-ring tissue. Higher stomatal conductances of the irrigated trees (0.22 vs. 0.08 mol m⁻² s⁻¹ on average) corresponded with ~2.0‰ lower average δ¹³C values, both measured and derived. Derived and measured δ¹³C values also indicated that needle growth, which occurs throughout the stressful summer was supported by carbon from concurrent, low rate assimilation. For Aleppo pine under semi-arid and irrigated conditions, the δ¹³C of tree-ring and needle material proved, in general, to be a reasonable indicator of integrated leaf gas-exchange properties.     

Determinants of isotopic coupling of CO2 and water vapour within a Quercus petraea forest canopy

Concentration and isotopic composition (δ¹³C and δ¹⁸O) of ambient CO₂ and water vapour were determined within a Quercus petraea canopy, Northumberland, UK. From continuous measurements made across a 36-h period from three heights within the forest canopy, we generated mixing lines (Keeling plots) for δ_a ¹³CO₂, δ_a C¹⁸O¹⁶O and δ_a H₂ ¹⁸O, to derive the isotopic composition of the signal being released from forest to atmosphere. These were compared directly with measurements of different respective pools within the forest system, i.e. δ¹³C of organic matter input for δ_a ¹³CO₂, δ¹⁸O of exchangeable water for δ_a C¹⁸O¹⁶O and transpired water vapour for δ_a H₂ ¹⁸O. [CO₂] and δ_a ¹³CO₂ showed strong coupling, where the released CO₂ was, on average, 4 per mil enriched compared to the organic matter of plant material in the system, suggesting either fractionation of organic material before eventual release as soil-respired CO₂, or temporal differences in ecosystem discrimination. δ_a C¹⁸O¹⁶O was less well coupled to [CO₂], probably due to the heterogeneity and transient nature of water pools (soil, leaf and moss) within the forest. Similarly, δ_a H₂ ¹⁸O was less coupled to [H₂O], again reflecting the transient nature of water transpired to the forest, seen as uncoupling during times of large changes in vapour pressure deficit. The δ¹⁸O of transpired water vapour, inferred from both mixing lines at the canopy scale and direct measurement at the leaf level, approximated that of source water, confirming that an isotopic steady state held for the forest integrated over the daily cycle. This demonstrates that isotopic coupling of CO₂ and water vapour within a forest canopy will depend on absolute differences in the isotopic composition of the respective pools involved in exchange and on the stability of each of these pools with time. Received: 21 March 1998 / Accepted: 10 December 1998     

Comparison of leaf water use efficiency of oak and sycamore in the canopy over two growing seasons

The seasonal trends in water use efficiency of sun and shade leaves of mature oak (Quercus robur) and sycamore (Acer pseudoplatanus) trees were assessed in the upper canopy of an English woodland. Intrinsic water use efficiency (net CO₂ assimilation rate/leaf conductance, A/g) was measured by gas exchange and inferred from C isotope discrimination (δ¹³C) methods. Shade leaves had consistently lower δ¹³C than sun leaves (by 1–2‰), the difference being larger in sycamore. Buds had distinct sun and shade isotopic signatures before bud break and received an influx of ¹³C-rich C before becoming net autotrophs. After leaf full expansion, δ¹³C declined by 1–2‰ gradually through the season, emphasising the importance of imported carbon in the interpretation of leaf δ¹³C values in perennial species. There was no significant difference between the two species in the value of intrinsic water use efficiency for either sun or shade leaves. For sun leaves, season-long A/g calculated from δ¹³C (72–78 μmol CO₂ [mol H₂O]⁻¹) was 10–16% higher than that obtained from gas exchange and in situ estimates of leaf boundary layer conductance. For shade leaves, the gas exchange–derived values were low, only 10–18% of the δ¹³C-derived values. This is ascribed to difficulties in obtaining a comprehensive sample of gas exchange measurements in the rapidly changing light environment.     

Diurnal variations of needle water isotopic ratios in two pine species

Diurnal fluctuations of leaf water isotope ratios (δ¹⁸O and δD) were measured for Jeffrey (Pinus jeffreyi Balf.) and lodgepole (Pinus contorta Douglas ex Louden) pine. Two trees per species were sampled every few hours on 15–16 October 2005 and 19–20 June 2006. Diurnal gas exchange was measured during the summer sampling. In fall 2005, leaf water δ¹⁸O ranged from 0.7 to 9.0‰, and leaf water δD ranged from −70 to −50‰. In summer 2006, leaf water δ¹⁸O ranged from 7.7 to 20.7‰, and leaf water δD ranged from −61 to −24‰. Diurnal variation of leaf water isotope values typically reached a maximum in early afternoon, began decreasing around midnight, and reached a minimum in mid-morning. Both periods showed a high degree of enrichment relative to source water, with leaf water–source water enrichments ranging up to 37.8‰ for δ¹⁸O, and up to 95‰ for δD. Leaf water enrichment varied by season with summer enrichment being greater than fall enrichment. A steady-state model (i.e., modified Craig–Gordon modeling) for leaf water isotope compositions did not provide a good fit to measured values of leaf water. In summer, a non-steady state model provided a better fit to the measured data than the steady-state model. Our findings demonstrate substantial leaf water enrichment above source water and diurnal variations in the isotopic composition of leaf water, which has application to understanding short-term variability of atmospheric gases (water vapor, CO₂, O₂), climate studies based on the isotopic composition of tree rings, and ecosystem water fluxes.