Drivers of radial growth and carbon isotope discrimination of bur oak (Quercus macrocarpa Michx.) across continental gradients in precipitation,vapour pressure deficit and irradiance

Tree‐ring characteristics are commonly used to reconstruct climate variables, but divergence from the assumption of a single biophysical control may reduce the accuracy of these reconstructions. Here, we present data from bur oaks (Quercus macrocarpa Michx.) sampled within and beyond the current species bioclimatic envelope to identify the primary environmental controls on ring‐width indices (RWIs) and carbon stable isotope discrimination (Δ¹³C) in tree‐ring cellulose. Variation in Δ¹³C and RWI was more strongly related to leaf‐to‐air vapour pressure deficit (VPD) at the centre and western edge of the range compared with the northern and wettest regions. Among regions, Δ¹³C of tree‐ring cellulose was closely predicted by VPD and light responses of canopy‐level Δ¹³C estimated using a model driven by eddy flux and meteorological measurements (R² = 0.96, P = 0.003). RWI and Δ¹³C were positively correlated in the drier regions, while they were negatively correlated in the wettest region. The strength and direction of the correlations scaled with regional VPD or the ratio of precipitation to evapotranspiration. Therefore, the correlation strength between RWI and Δ¹³C may be used to infer past wetness or aridity from paleo wood by determining the degree to which carbon gain and growth have been more limited by moisture or light.     

The trophic structure of bark-living oribatid mite communities analysed with stable isotopes (15N, 13C) indicates strong niche differentiation

The aim of the present study was to identify food sources of bark-living oribatid mites to investigate if trophic niche differentiation contributes to the diversity of bark living Oribatida. We measured the natural variation in stable isotope ratios (¹⁵N/¹⁴N, ¹³C/¹²C) in oribatid mites from the bark of oak (Quercus robur), beech (Fagus sylvatica), spruce (Picea abies) and pine (Pinus sylvestris) trees and their potential food sources, i.e., the covering vegetation of the bark (bryophytes, lichens, algae, fungi). As a baseline for calibration the stable isotope signatures of the bark of the four tree species were measured and set to zero. Oribatid mite stable isotope ratios spanned over a range of about 13 δ units for ¹⁵N and about 7 δ units for ¹³C suggesting that they span over about three trophic levels. Different stable isotope signatures indicate that bark living oribatid mites feed on different food sources, i.e., occupy distinct trophic niches. After calibration stable isotope signatures of respective oribatid mite species of the four tree species were similar indicating close association of oribatid mites with the corticolous cover as food source. Overall, the results support the hypothesis that trophic niche differentiation of bark living oribatid mites contributes to the high diversity of the group.     

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.     

Modulation of leaf attributes and water use efficiency in Quercus suber along a rainfall gradient

The aim of our study was to assess the intraspecific variation of a range of leaf attributes and carbon isotope discrimination (Δ), in Quercus suber, along an 800 mm rainfall gradient in Portugal. We measured specific leaf area (SLA), leaf thickness (LT) and density, and used leaf carbon isotope content as an integrated record of water use efficiency. The values of SLA of Q. suber were strongly, positively correlated with rainfall, decreasing as rainfall decreased. This reduction was mainly driven by LT, which was tightly correlated with SLA (R = −0.80, P < 0.001), and steadily increased with declining rainfall. The significant increase in carbon isotope discrimination towards the wetter end of the gradient, with a difference of 4.2‰ in Δ between the two extremes, suggests a strong adjustment of leaf gas exchange to water availability. Leaf changes associated with precipitation in Q. suber thus seem to influence water economy, since reduction of SLA with the increase of LT with aridity improves water use efficiency. These data suggest that this evergreen tree species relies on its foliage plasticity and physiology to overcome water shortage.     

Comparison of seasonal variations in water-use efficiency calculated from the carbon isotope composition of tree rings and flux data in a temperate forest

Tree-ring δ(13) C is often interpreted in terms of intrinsic water-use efficiency (WUE) using a carbon isotope discrimination model established at the leaf level. We examined whether intra-ring δ(13) C could be used to assess variations in intrinsic WUE (W(g), the ratio of carbon assimilation and stomatal conductance to water) and variations in ecosystem WUE (W(t) , the ratio of C assimilation and transpiration) at a seasonal scale. Intra-ring δ(13) C was measured in 30- to 60-μm-thick slices in eight oak trees (Quercus petraea). Canopy W(g) was simulated using a physiologically process-based model. High between-tree variability was observed in the seasonal variations of intra-ring δ(13) C. Six trees showed significant positive correlations between W(g) calculated from intra-ring δ(13) C and canopy W(g) averaged over several days during latewood formation. These results suggest that latewood is a seasonal recorder of W(g) trends, with a temporal lag corresponding to the mixing time of sugars in the phloem. These six trees also showed significant negative correlations between photosynthetic discrimination Δ calculated from intra-ring δ(13) C, and ecosystem W(t), during latewood formation. Despite the observed between-tree variability, these results indicate that intra-ring δ(13) C can be used to access seasonal variations in past W(t).     

Comparative nutrient economy,stable isotopes,and related adaptive traits in <Emphasis Type="Italic">Picea rubens</Emphasis>, <Emphasis Type="Italic">Picea mariana</Emphasis>, and their hybrids

Nutrient- and water economy-related traits in plants have significant implications for growth and fitness. We explored, examined, and compared nutrient concentrations, use efficiencies, assimilation, and informative isotopic elements in a seedling provenance experiment, and in seedling and mature tree controlled-cross hybrid experiments of red spruce (RS) (Picea rubens Sarg.) and black spruce (BS) (P. mariana (Mill.) BSP). Provenance experiment results showed RS had consistently lower carbon (C) and nitrogen (N) concentrations, and N assimilation ratio (NAR), but higher N-use efficiency (NUE), C:N ratio, water-use efficiency (WUE), and needle calcium (Ca) and magnesium (Mg) concentrations than BS. The hybrid seedling experiment showed similar results and additive inheritance for needle N, C:N ratio, NAR, Ca, and Mg, evident by a near-linear progression from one species to the other. Within both species, seedling height showed a negative relationship with needle N and a positive relationship with NUE. However, across hybrid indices, seedling height showed a positive relationship with needle N and a negative relationship with NUE. Also across hybrid indices, seedling height showed a negative relationship to Ca and C:N, and a positive relationship with NAR and ¹³C discrimination (without hybrid 25). Mature tree hybrid experiment results were similar to those of the seedling experiment, but with a dampening of differences caused by low nutrient availability and possibly age effects. The similarity was not true for ¹³C discrimination as mature tree height showed a strong negative relationship to ¹³C discrimination, indicating that BS had greater WUE. The reversal is most probably caused by the large difference in water availability.     

Variations in Environmental Signals in Tree-Ring Indices in Trees with Different Growth Potential

We analysed two groups of Quercus robur trees, growing at nearby plots with different micro-location condition (W-wet and D-dry) in the floodplain Krakovo forest, Slovenia. In the study we compared the growth response of two different tree groups to environmental variables, the potential signal stored in earlywood (EW) structure and the potential difference of the information stored in carbon isotope discrimination of EW and latewood (LW). For that purpose EW and LW widths and carbon isotope discrimination for the period 1970–2008 AD were measured. EW and LW widths were measured on stained microscopic slides and chronologies were standardised using the ARSTAN program. α-cellulose was extracted from pooled EW and LW samples and homogenized samples were further analysed using an elemental analyser and IRMS. We discovered that W oaks grew significantly better over the whole analysed period. The difference between D and W oaks was significant in all analysed variables with the exception of stable carbon isotope discrimination in latewood. In W oaks, latewood widths correlated with summer (June to August) climatic variables, while carbon isotope discrimination was more connected to River Krka flow during the summer. EW discrimination correlated with summer and autumn River Krka flow of the previous year, while latewood discrimination correlated with flow during the current year. In the case of D oaks, the environmental signal appears to be vague, probably due to less favourable growth conditions resulting in markedly reduced increments. Our study revealed important differences in responses to environmental factors between the two oak groups of different physiological conditions that are preconditioned by environmental stress. Environmental information stored in tree-ring features may vary, even within the same forest stand, and largely depends on the micro-environment. Our analysis confirmed our assumptions that separate EW and LW analysis of widths and carbon isotope discrimination provides complementary information in Q. robur dendroecology.     

Leaf carbon isotope ratio and water use efficiency of urban roadside trees in summer in Kyoto city

The effect of summer climate on leaf carbon isotope composition (δ¹³C) of the major roadside tree species Prunus × yedoensis (P. yedoensis) was investigated in Kyoto city, Japan, to explore the implications for alterations in urban environments. Temperature and the vapor pressure deficit were higher at sites of higher traffic volumes, possibly affected by a heat island effect. The leaf δ¹³C of P. yedoensis trees was affected strongly by leaf carbon isotope discrimination (Δ), with much less effect of δ¹³C on atmospheric CO₂. Leaf Δ values in the summer were smaller at sites of higher traffic volumes with high atmospheric temperatures, suggesting a higher long-term water use efficiency (WUE) at these sites. Gas exchange measurements of P. yedoensis leaves indeed suggested a higher intrinsic WUE at sites of higher traffic volumes with high atmospheric temperatures. These results suggest that leaf Δ is related to the response of WUE to summer climates, and that leaf δ¹³C in urban areas is a useful tracer for understanding the influences of urban environments on plant photosynthetic processes.     

Stable carbon isotopes and the metabolism of the terrestrial Devonian organism Spongiophyton

Devonian fossils of Spongiophyton have been identified as a terrestrial evolutionary intermediate between algae and vascular land plants on the basis of their dichotomously branched, tubular morphology, thick cuticles, and the scattered distribution of surface pores. Our understanding of their physiology is, however, severely limited, but may be increased through the use of stable carbon isotope measurements. One such study led to the hypothesis that Spongiophyton carried a carbon isotope (δ¹³C) signature characteristic of lichens ( Jahren et al., 2003 Geology 31 , 99–102). Here, we outline three difficulties with accepting this idea and report independent isotopic measurements of Spongiophyton fossils from Canada and Ghana. Our results show that the isotopic discrimination of analogous tissues of Spongiophyton, extant lichens, liverworts and mosses are statistically indistinguishable. We suggest therefore that claims to have definitively identified lichen metabolism are premature, and cannot be sustained.     

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.     

Disentangling the effects of acidic air pollution,atmospheric CO2, and climate change on recent growth of red spruce trees in the Central Appalachian Mountains

In the 45 years after legislation of the Clean Air Act, there has been tremendous progress in reducing acidic air pollutants in the eastern United States, yet limited evidence exists that cleaner air has improved forest health. Here, we investigate the influence of recent environmental changes on the growth and physiology of red spruce (Picea rubens Sarg.) trees, a key indicator species of forest health, spanning three locations along a 100 km transect in the Central Appalachian Mountains. We incorporated a multiproxy approach using 75‐year tree ring chronologies of basal tree growth, carbon isotope discrimination (?¹³C, a proxy for leaf gas exchange), and δ¹⁵N (a proxy for ecosystem N status) to examine tree and ecosystem level responses to environmental change. Results reveal the two most important factors driving increased tree growth since ca. 1989 are reductions in acidic sulfur pollution and increases in atmospheric CO₂, while reductions in pollutant emissions of NO_x and warmer springs played smaller, but significant roles. Tree ring ?¹³C signatures increased significantly since 1989, concurrently with significant declines in tree ring δ¹⁵N signatures. These isotope chronologies provide strong evidence that simultaneous changes in C and N cycling, including greater photosynthesis and stomatal conductance of trees and increases in ecosystem N retention, were related to recent increases in red spruce tree growth and are consequential to ecosystem recovery from acidic pollution. Intrinsic water use efficiency (iWUE) of the red spruce trees increased by ~51% across the 75‐year chronology, and was driven by changes in atmospheric CO₂ and acid pollution, but iWUE was not linked to recent increases in tree growth. This study documents the complex environmental interactions that have contributed to the recovery of red spruce forest ecosystems from pervasive acidic air pollution beginning in 1989, about 15 years after acidic pollutants started to decline in the United States.     

The dendroclimatic value of oak stable isotopes

Tree-ring stable oxygen and carbon isotope ratios (δ¹⁸O and δ¹³C) are an important archive for climate reconstructions. However, it remains unclear whether the polyvinyl acetate emulsion, often used for the preservation and fixation of wood samples, influences δ¹⁸O and δ¹³C signals. Further uncertainties are associated with the possible effects of geographical origin and cambial age of historical samples. Here, we present annually-resolved and absolutely-dated δ¹⁸O and δ¹³C measurements of 21 living oaks (Quercus robur and Q. petraea) from the Czech Republic. We find that the δ¹⁸O and δ¹³C signals in the extracted alpha-cellulose are not affected by polyvinyl acetate treatment. Covering the entire 20^th century and reaching until 2018 CE, our dataset reveals spatial and temporal coherency within and between the individual δ¹⁸O and δ¹³C chronologies of different oak species, sample locations, and tree ages. Highly significant (p < 0.01) Pearson’s correlation coefficients of the site-specific δ¹³C and δ¹⁸O chronologies range from 0.48–0.77 and 0.36–0.56, respectively. The isotopic inter-series correlations of Q. robur and Q. petraea from the same site are 0.75 and 0.43 for the mean δ¹³C and δ¹⁸O values, respectively. Significant (p < 0.01) correlations of 0.49 and 0.84 are found for δ¹³C and δ¹⁸O, respectively, when all measurements from all sampling locations and tree ages are included. Our study shows that non-pooled oak δ¹⁸O and δ¹³C measurements from both species, different locations, and diverse tree ages can be combined into robust isotopic chronologies for climate reconstructions.     

Concentration and {sigma}13C of leaf carbohydrates in relation to gas exchange in Quercus robur under elevated CO2 and drought

The variations of leaf carbohydrate concentration, carbon isotopediscrimination () of leaf soluble carbohydrate, gas-exchangeand growth during a soil drying cycle under 350 and 700 µmolmol^-1 CO₂ concentrations ([CO₂]) inQuercus robur seedlings wereanalysed. In well-watered conditions, a doubling of [CO₂] causedan increase of CO₂ assimilation rate (A) ( +47%) and a decreaseof stomatal conductance for water vapour (g) (–25%),anddoubled the intrinsic water-use efficiency (A/g). The valuesof A were not affected by elevated [CO₂] which was consistentwith the 2-fold increase of A/g. Elevated [CO₂ also significantlyincreased sucrose and starch leaf concentrations as well asaerial growth and plant dry weight. The stimulating effect ofCO₂ enrichment on A and A/g was maintained in moderate droughtconditions, but disappeared in the most severe drought conditions.Drought induced an increase of hexose concentrations in both[CO₂], but this effect was more pronounced under elevated [CO₂],which may contribute to increase osmoregulation. From the onsetof drought, starch was depleted in both [CO₂]. Carbon isotopediscrimination decreased in response to drought, which correspondedto an increase in A/g according to the two-step model of isotopicdiscrimination. In contrast, the A/g values derived from instantaneousleaf gas-exchange measurements decreased along the drying cycle.The discrepancy observed between the two independent estimatesof water-use efficiency is discussed in terms of time-scaleintegration. The results obtained with the isotopic approachusing soluble carbohydrate suggest a predominant stomatal limitationof CO2 assimilation in response to drought. Soil drying cycle,elevated CO₂, leaf gas-exchange, leaf carbohydrate concentrations,carbon isotope discrimination, growth, Quercus robur. Key words: soil drying cycle, elevated CO₂, leaf gas-exchange, leaf carbohydrate concentrations, carbon isotope discrimination, growth, Quercus robur     

Fractionation of Carbon Isotopes during Biogenesis of Atmospheric Isoprene

The stable carbon isotope composition of isoprene emitted from leaves of red oak (Quercus rubra L.) was measured. Isoprene was depleted in ¹³C relative to carbon recently fixed by photosynthesis. The difference in isotope composition between recently fixed carbon and emitted isoprene was independent of the isotopic composition of the source CO₂. β-Carotene, an isoprenoid plant constituent, was depleted in ¹³C relative to whole leaf carbon to the same degree as isoprene, but fatty acids were more depleted. Isoprene emitted from leaves fed abscisic acid was much less depleted in ¹³C than was isoprene emitted from unstressed leaves. We conclude that isoprene is made from an isoprenoid precursor that is derived from acetyl-CoA made from recent photosynthate. The carbon isotope composition of isoprene in the atmosphere is likely to be slightly more negative (less ¹³C) than C₃ plant material but when plants are stressed the isotopic composition could vary.     

Tracing site‐specific isotopic signatures along a Blue Tit Cyanistes caeruleus food chain

Food chains culminating with temperate insectivorous passerines are well described, yet whether trophic webs can be site‐specific remains a largely unexplored question. In the case of site‐ or habitat‐specificity of food webs, stable isotope signatures of bird feathers may enable assignment of unmarked individuals to a site or a habitat of origin. We address this question in landscapes that include contrasting forest habitat patches with either deciduous Downy Oak Quercus humilis or evergreen Holm Oak Quercus ilex as dominant tree species. First, we examine the spatial variation across habitats and sites in the stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) along the oak leaf–Tortrix moth Tortrix viridana caterpillar–Blue Tit Cyanistes caeruleus food chain. Secondly, we assess whether the isotopic signatures allow for correct assignment of individual birds to their site or habitat of origin. At the scale of the landscape, stable isotope values enabled identification of the different components of the Blue Tit food chain: from oak leaves to Blue Tit nestlings and yearling birds. However, isotopic signatures were site‐specific (i.e. geographical) more than habitat‐specific (i.e. deciduous vs. evergreen oaks). Discriminant analyses correctly assigned 85% of nestlings and 83% of resident yearling birds, indicating a pronounced effect of site on Blue Tit feather isotopic signatures. We thereby demonstrate that isotopes reflect a stronger association of locally born birds to the local features of their habitat than that of un‐ringed yearling birds, whose plumage may have grown while in a wider geographical area. This study provides evidence of site‐specific isotopic signatures from oak leaves to Blue Tit feathers at a fine spatial scale.     

Ectomycorrhizal fungal communities of native and non-native Pinus and Quercus species in a common garden of 35-year-old trees

Non-native tree species have been widely planted or have become naturalized in most forested landscapes. It is not clear if native trees species collectively differ in ectomycorrhizal fungal (EMF) diversity and communities from that of non-native tree species. Alternatively, EMF species community similarity may be more determined by host plant phylogeny than by whether the plant is native or non-native. We examined these unknowns by comparing two genera, native and non-native Quercus robur and Quercus rubra and native and non-native Pinus sylvestris and Pinus nigra in a 35-year-old common garden in Poland. Using molecular and morphological approaches, we identified EMF species from ectomycorrhizal root tips and sporocarps collected in the monoculture tree plots. A total of 69 EMF species were found, with 38 species collected only as sporocarps, 18 only as ectomycorrhizas, and 13 both as ectomycorrhizas and sporocarps. The EMF species observed were all native and commonly associated with a Holarctic range in distribution. We found that native Q. robur had ca. 120% higher total EMF species richness than the non-native Q. rubra, while native P. sylvestris had ca. 25% lower total EMF species richness than non-native P. nigra. Thus, across genera, there was no evidence that native species have higher EMF species diversity than exotic species. In addition, we found a higher similarity in EMF communities between the two Pinus species than between the two Quercus species. These results support the naturalization of non-native trees by means of mutualistic associations with cosmopolitan and novel fungi.     

Leaf gas exchange and carbon isotope composition responses to drought in a drought-avoiding (Pinus pinaster) and a drought-tolerant (Quercus petraea) species under present and elevated atmospheric CO2 concentrations

The responses of predawn leaf water potential (φ_wp), leaf conductance to water vapour diffusion (g), CO₂ assimilation rate (A) and carbon isotope competition (δ¹³C) to a soil drying cycle were assessed in Pinus pinaster, a drought-avoiding species with high stomatal sensitivity to drought, and Quercus petraea, a drought-tolerant species with lower stomatal sensitivity to drought, under present (350 μmol^?1) and elevated (700 μmol^?1) atmospheric CO₂ concentrations ([CO₂]). In P. pinaster, decreasing A in response to drought was associated with increasing plant intrinsic water use efficiency (A/g) and with decreasing calculated intercellular [CO₂] (C₁), suggesting a stomatal limitation of A. In contrast, in Q. petraea, A/g declined and C¹ increased during the drying cycle, which suggests a non-stomatal origin for the decrease in A. In P. pinaster, a negative relationship was observed between the gas exchange-derived values of C_i/C_a and δ¹³C, which conforms to the classical two-step carbon isotope discrimination model. In Q. petraea, the relationship between C₁/C_a and δ¹³C was positive. Possible causes of this discrepancy are discussed. Lower g values were observed under elevated [CO₂] than under present [CO₂] in Q. petraea, whereas g was unaffected in P. pinaster. A stimulation of A by elevated [CO₂] was found in P. pinaster but not in Q. petraea. In both species, A/g was markedly higher under elevated than under present [CO₂]. Whether the differences in the g response to elevated [CO₂] found here can be generalized to other drought-avoiding and non-avoiding species remains to be assessed.     

Stomatal conductance, photosynthesis and respiration of temperate deciduous tree seedlings grown outdoors at an elevated concentration of carbon dioxide

Seedlings of temperate deciduous tree species were grown outdoors at ambient and at an elevated concentration of carbon dioxide to examine how aspects of their gas exchange would be altered by growth at elevated carbon dioxide concentration. Leaf conductances to water vapour and net carbon dioxide exchange rates were determined periodically near midday. Whole-plant carbon dioxide efflux rates in darkness were also determined. The stomatal conductance of leaves of plants grown and measured at 700 cm³ m^?3 carbon dioxide did not differ from that of plants grown and measured at 350 cm³ m^?3 in Malus domestica, Quercus prinus and Quercus robur at any measurement time. In Acer saccharinum, lower conductances occurred for plants grown and measured at elevated carbon dioxide concentration only at measurement temperatures above 33°C. Photo-synthetic adjustment to elevated carbon dioxide concentration was evident only in Q. robur. All species examined had lower rates of dark respiration per unit of mass when grown and measured at elevated carbon dioxide concentration.     

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.