Ontogeny and leaf gas exchange mediate the carbon isotopic signature of herbaceous plants

Values (Δ(i)) predicted by a simplified photosynthetic discrimination model, based only on diffusion through air followed by carboxylation, are often used to infer ecological conditions from the 13C signature of plant organs (δ13C(p)). Recent studies showed that additional isotope discrimination (d that includes mesophyll conductance, photorespiration and day respiration, and post-carboxylation discrimination) can strongly affect δ13C(p); however, little is known about its variability during plant ontogeny for different species. Effect of ontogeny on leaf gas exchange rates, Δ(i) , observed discrimination (Δ(p)) and d in leaf, phloem and root of seven herbaceous species at three ontogenetic stages were investigated under controlled conditions. Functional group identity and ontogeny significantly affected Δ(i) and Δ(p). However, predicted Δ(i) did not match Δ(p). d, strongly affected by functional group identity and ontogeny, varied by up to 14 ‰. d scaled tightly with stomatal conductance, suggesting complex controls including changes in mesophyll conductance. The magnitude of the changes in δ13C(p) due to ontogeny was similar to that due to environmental factors reported in other studies. d and ontogeny should, therefore, be considered in ecosystem studies, integrated in ecosystem models using δ13C(p) and limit the applicability of δ13C(leaf) as a proxy for water-use efficiency in herbaceous plants.     

Carbon isotopic composition and oxygen isotopic enrichment in phloem and total leaf organic matter of European beech (Fagus sylvatica L.) along a climate gradient

This study investigated the influence of climate on the carbon isotopic composition (sigma13C) and oxygen isotopic enrichment (delta18O) above the source water of different organic matter pools in European beech. In July and September 2002, sigma13C and delta18O were determined in phloem carbohydrates and in bulk foliage of adult beech trees along a transect from central Germany to southern France, where beech reaches its southernmost distributional limit. The data were related to meteorological and physiological parameters. The climate along the transect stretches from temperate [subcontinental (SC)] to submediterranean (SM). Both sigma13Cleaf and delta18Oleaf were representative of site-specific long-term environmental conditions. sigma13C of leaves collected in September was indicative of stomatal conductance, vapour pressure deficit (VPD) and radiation availability of the current growing season. delta18O was mainly correlated to mean growing season relative humidity (RH) and VPD. In contrast to the leaves, sigma13Cphloem varied considerably between July and September and was well correlated with canopy stomatal conductance (Gs) in a 2 d integral prior to phloem sampling. The relationship between sigma13C and delta18O in both leaves and phloem sap points, however, to a combined influence of stomatal conductance and photosynthetic capacity on the variation of sigma13C along the transect. delta18Ophloem could be described by applying a model that included 18O fractionation associated with water exchange between the leaf and the atmosphere and with the production of organic matter. Hence, isotope signatures can be used as effective tools to assess the water balance of beech, and thus, help predict the effects of climatic change on one of the ecologically and economically most important tree species in Central Europe.     

Stand structure modulates the long-term vulnerability of Pinus halepensis to climatic drought in a semiarid Mediterranean ecosystem

We investigated whether stand structure modulates the long-term physiological performance and growth of Pinus halepensis Mill. in a semiarid Mediterranean ecosystem. Tree radial growth and carbon and oxygen stable isotope composition of latewood (δ(13)C(LW) and δ(18)O(LW), respectively) from 1967 to 2007 were measured in P. halepensis trees from two sharply contrasting stand types: open woodlands with widely scattered trees versus dense afforested stands. In both stand types, tree radial growth, δ(13)C(LW) and δ(18)O(LW) were strongly correlated with annual rainfall, thus indicating that tree performance in this semiarid environment is largely determined by inter-annual changes in water availability. However, trees in dense afforested stands showed consistently higher δ(18)O(LW) and similar δ(13)C(LW) values compared with those in neighbouring open woodlands, indicating lower stomatal conductance and photosynthesis rates in the former, but little difference in water use efficiency between stand types. Trees in dense afforested stands were more water stressed and showed lower radial growth, overall suggesting greater vulnerability to drought and climate aridification compared with trees in open woodlands. In this semiarid ecosystem, the negative impacts of intense inter-tree competition for water on P. halepensis performance clearly outweigh potential benefits derived from enhanced infiltration and reduced run-off losses in dense afforested stands.     

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).     

Combining delta 13 C and delta 18 O analyses to unravel competition, CO2 and O3 effects on the physiological performance of different-aged trees

Combined delta(13)C and delta(18)O analyses of leaf material were used to infer changes in photosynthetic capacity (A(max)) and stomatal conductance (g(l)) in Fagus sylvatica and Picea abies trees growing under natural and controlled conditions. Correlation between g(l) and delta(18)O in leaf cellulose (delta(18)O(cel)) allowed us to apply a semi-quantitative model to infer g(l) from delta(18)O(cel) and also interpret variation in delta(13)C as reflecting variation in A(max). Extraction of leaf cellulose was necessary, because delta(18)O from leaf organic matter (delta(18)O(LOM)) and delta(18)O(cel) was not reliably correlated. In juvenile trees, the model predicted elevated carbon dioxide (CO(2)) to reduce A(max) in both species, whereas ozone (O(3)) only affected beech by reducing CO(2) uptake via lowered g(l). In adult trees, A(max) declined with decreasing light level as g(l) was unchanged. O(3) did not significantly affect isotopic signatures in leaves of adult trees, reflecting the higher O(3) susceptibility of juvenile trees under controlled conditions. The isotopic analysis compared favourably to the performance of leaf gas exchange, underlining that the semi-quantitative model approach provides a robust way to gather time-integrated information on photosynthetic performance of trees under multi-faced ecological scenarios, in particular when information needed for quantitative modelling is only scarcely available.     

不同土壤含水量下侧柏幼树叶片水分利用效率

对植物水分利用效率的研究,可以揭示植物的内在耗水机制,为区域森林生态系统经营与维护提供依据.本研究以侧柏幼树为研究对象,通过室内控制试验设置不同的土壤水分梯度,分别用气体交换法和稳定同位素法对其不同土壤含水量条件下的瞬时水分利用效率(WUE_gs)和短期水分利用效率(WUE_cp)进行研究.结果表明:受气孔导度(g_s)的影响,叶片净光合速率(P_n)和蒸腾速率(T_r)随土壤含水量的增加呈现相同的变化趋势,均在土壤含水量为70%～80%田间持水量(FC)时达到最大值;叶片WUE_gs则在土壤含水量最低(35%～45% FC)时达到最大值(7.26 mmol·m^-2·s^-1).叶片可溶性糖、枝条韧皮部渗出液的δ¹³C都在土壤含水量最低(35%～45% FC)条件下达到最大值,且叶片可溶性糖的δ¹³C明显高于枝条韧皮部渗出液的δ¹³C,未产生明显分馏;而叶片WUE_cp也在土壤含水量最低(35%～45% FC)时达到最大值(7.26 mmol·m^-2·s^-1).相同条件下,叶片WUE_gs和WUE_cp存在一定差异(平均相差0.52 mmol·m^-2·s^-1),WUE_gs时空变异性较大,而WUE_cp更具有代表性.侧柏幼树通过降低生理生态活动和提高水分利用效率来适应干旱的土壤条件.     

Century‐long apparent decrease in intrinsic water‐use efficiency with no evidence of progressive nutrient limitation in African tropical forests

Forests exhibit leaf‐ and ecosystem‐level responses to environmental changes. Specifically, rising carbon dioxide (CO₂) levels over the past century are expected to have increased the intrinsic water‐use efficiency (iWUE) of tropical trees while the ecosystem is gradually pushed into progressive nutrient limitation. Due to the long‐term character of these changes, however, observational datasets to validate both paradigms are limited in space and time. In this study, we used a unique herbarium record to go back nearly a century and show that despite the rise in CO₂ concentrations, iWUE has decreased in central African tropical trees in the Congo Basin. Although we find evidence that points to leaf‐level adaptation to increasing CO₂—that is, increasing photosynthesis‐related nutrients and decreasing maximum stomatal conductance, a decrease in leaf δ¹³C clearly indicates a decreasing iWUE over time. Additionally, the stoichiometric carbon to nitrogen and nitrogen to phosphorus ratios in the leaves show no sign of progressive nutrient limitation as they have remained constant since 1938, which suggests that nutrients have not increasingly limited productivity in this biome. Altogether, the data suggest that other environmental factors, such as increasing temperature, might have negatively affected net photosynthesis and consequently downregulated the iWUE. Results from this study reveal that the second largest tropical forest on Earth has responded differently to recent environmental changes than expected, highlighting the need for further on‐ground monitoring in the Congo Basin.     

Relationships of growth,stable carbon isotope composition and anatomical properties of leaf and xylem in seven mulberry cultivars: a hint towards drought tolerance

• The fast growth of mulberry depends on high water consumption, but considerable variations in drought tolerance exist across different cultivars. Physiological and anatomical mechanisms are important to plant survival under drought. However, few research efforts have been made to reveal the relationships of these two aspects in relation to drought tolerance.
• In this study, growth rates, leaf functional physiology and anatomical characteristics of leaf and xylem of 1‐year‐old saplings of seven mulberry cultivars at a common garden were compared. Their relationships were also explored.
• Growth, leaf physiology and anatomy were significantly different among the tested cultivars. Foliar stable carbon isotope composition (δ¹³C) was negatively correlated with growth rates, and closely related to several leaf and xylem anatomical traits. Particularly, leaf thickness, predicted hydraulic conductivity and vessel element length jointly contributed 77% of the variability in δ¹³C. Cultivar Wupu had small stomata, intermediate leaf thickness, the smallest hydraulically weighted vessel diameter and highest vessel number, and higher δ¹³C; Yunguo1 had high abaxial stomatal density, low specific leaf area, moderate hydraulic conductivity and δ¹³C; these are beneficial features to reduce leaf water loss and drought‐induced xylem embolism in arid areas. Cultivar Liaolu11 had contrasting physiological and anatomical traits compared with the previous two cultivars, suggesting that it might be sensitive to drought.
• Our findings indicate that growth and δ¹³C are closely associated with both leaf and xylem anatomical characteristics in mulberry, which provides fundamental information to assist evaluation of drought tolerance in mulberry cultivars and in other woody trees.

     

Carbon and oxygen isotope composition of organic compounds in the phloem sap provides a short-term measure for stomatal conductance of European beech (Fagus sylvatica L.)

At eight different dates during the 2000 growing season, δ¹³C and δ¹⁸O were determined in the phloem of adult beech trees growing in natural beech stands in south‐west Germany differing in stand density and local climate. In addition, stand transpiration, precipitation, photosynthetic active radiation, relative air humidity, water pressure deficit of the air, air and soil temperature, soil water potential, and sugar concentration of the phloem sap were determined directly and evaporation and canopy stomatal conductance were modelled. All parameters were related to δ¹³C. The study aimed to identify the time integral within which the δ¹³C of organic compounds transported in the phloem is an indicative measure of these environmental influences. δ¹³C of soluble carbon transported in the phloem was well correlated with mean stomatal conductance in a two‐day integral prior to phloem sampling but did not depend on either light intensity or soil water availability. A strong positive relationship between δ¹³C and δ¹⁸O pointed to observed variation in δ¹³C of phloem sap being a result of variation in stomatal conductance. Bulk leaf δ¹³C was a poor indicator of changes in environmental conditions during the growing season. From these results we conclude that the analysis of δ¹³C in soluble carbon transported in the phloem is a reliable indicator of short‐term changes in C_i/C_a. In contrast, the δ¹³C of structural carbon in beech foliage represents an integration of a range of factors that mask short‐term influences responsible for C_i/C_a.     

Compound-specific differences in (13)C of soluble carbohydrates in leaves and phloem of 6-month-old Eucalyptus globulus (Labill)

Movement of photoassimilates from leaves to phloem is an important step for the flux of carbon through plants. Fractionation of carbon isotopes during this process may influence their abundance in heterotrophic tissues. We subjected Eucalyptus globulus to 20, 25 and 28 °C ambient growth temperatures and measured compound-specific δ(13)C of carbohydrates obtained from leaves and bled phloem sap. We compared δ(13)C of sucrose and raffinose obtained from leaf or phloem and of total leaf soluble carbon, with modelled values predicted by leaf gas exchange. Changes in δ(13)C of sucrose and raffinose obtained from either leaves or phloem sap were more tightly coupled to changes in c(i)/c(a) than was δ(13)C of leaf soluble carbon. At 25 and 28 °C, sucrose and raffinose were enriched in (13)C compared to leaf soluble carbon and predicted values - irrespective of tissue type. Phloem sucrose was depleted and raffinose enriched in (13)C compared to leaf extracts. Intermolecular and tissue-specific δ(13)C reveal that multiple systematic factors influence (13)C composition during export to phloem. Predicting sensitivity of these factors to changes in plant physiological status will improve our ability to infer plant function at a range of temporal and spatial scales.     

Leaf water 18O and 2H enrichment along vertical canopy profiles in a broadleaved and a conifer forest tree

Distinguishing meteorological and plant‐mediated drivers of leaf water isotopic enrichment is prerequisite for ecological interpretations of stable hydrogen and oxygen isotopes in plant tissue. We measured input and leaf water δ²H and δ¹⁸O as well as micrometeorological and leaf morpho‐physiological variables along a vertical gradient in a mature angiosperm (European beech) and gymnosperm (Douglas fir) tree. We used these variables and different enrichment models to quantify the influence of Péclet and non‐steady state effects and of the biophysical drivers on leaf water enrichment. The two‐pool model accurately described the diurnal variation of leaf water enrichment. The estimated unenriched water fraction was linked to leaf dry matter content across the canopy heights. Non‐steady state effects and reduced stomatal conductance caused a higher enrichment of Douglas fir compared to beech leaf water. A dynamic effect analyses revealed that the light‐induced vertical gradients of stomatal conductance and leaf temperature outbalanced each other in their effects on evaporative enrichment. We conclude that neither vertical canopy gradients nor the Péclet effect is important for estimates and interpretation of isotopic leaf water enrichment in hypostomatous trees. Contrarily, species‐specific non‐steady state effects and leaf temperatures as well as the water vapour isotope composition need careful consideration.     

Stomatal activity within the crowns of tall deciduous trees under forest conditions

The variation in stomatal activity within the crowns ofAcer campestre, Carpinus betulus andQuercus cerris was measured by vapour exchange porometer on several summer days in an oak-hornbeam forest, in SW Slovakia, Czechoslovakia. Variation resulted from crown position in the forest stand and from leaf position within the canopy. The highest stomatal conductance was in sunlit sun leaves in the upper part of the canopy. Stomatal conductance decreased with increasing depth in the canopy. The steepest decrease was in the upper canopy, in the intermediate zone between fully sunlit and fully shaded leaves, and was caused by the decline in leaf irradiance and in stomatal density. In codominant trees, the conductance in shade leaves at the base of the crown was significantly lower than in the sun leaves at the top of the crown. In a dominant tree,Q. cerris, the differences in stomatal conductance were small and most frequently insignificant. Variation in incident light also determined the diurnal variation of stomatal conductance with respect to crown aspect. Differences between sun leaves on the east and west facing aspects of the overstory crown ofQ. cerris were demonstrated for several days.     

Isotopes reveal contrasting water use strategies among coexisting plant species in a Mediterranean ecosystem

Variation in the stable carbon and oxygen isotope composition (δ(13) C, Δ(18) O) of co-occurring plant species may reflect the functional diversity of water use strategies present in natural plant communities. We investigated the patterns of water use among 10 coexisting plant species representing diverse taxonomic groups and life forms in semiarid southeast Spain by measuring their leaf δ(13) C and Δ(18) O, the oxygen isotope ratio of stem water and leaf gas exchange rates. Across species, Δ(18) O was tightly negatively correlated with stomatal conductance (g(s) ), whereas δ(13) C was positively correlated with intrinsic water use efficiency (WUE(i) ). Broad interspecific variation in Δ(18) O, δ(13) C and WUE(i) was largely determined by differences in g(s) , as indicated by a strong positive correlation between leaf δ(13) C and Δ(18) O across species The 10 co-occurring species segregated along a continuous ecophysiological gradient defined by their leaf δ(13) C and Δ(18) O, thus revealing a wide spectrum of stomatal regulation intensity and contrasting water use strategies ranging from 'profligate/opportunistic' (high g(s) , low WUE(i) ) to 'conservative' (low g(s) , high WUE(i) ). Coexisting species maintained their relative isotopic rankings in 2?yr with contrasting rainfall, suggesting the existence of species-specific 'isotopic niches' that reflect ecophysiological niche segregation in dryland plant communities.     

Carbon isotope discrimination and stomatal responses of mature Pinus sylvestris L. trees exposed in situ for three years to elevated CO2 and temperature

The Climate Change Experiment (CLIMEX) is a unique large scale facility in which an entire undisturbed catchment of boreal vegetation has been exposed to elevated CO₂ (560 ppm) and temperature (+3°C summer, +5°C winter) for the past three years with all the soil-plant-atmosphere linkages intact. Here, carbon isotope composition and stomatal density have been analysed from sequential year classes of needles of mature Scots pine trees (Pinus sylvestris L.) to investigate the response of time-integrated water-use efficiency (UWE) and stomatal density to CO₂ enrichment and climate change. Carbon isotope discrimination decreased and WUE increased in cohorts of needles developing under increased CO₂ and temperature, compared to needles on the same trees developing in pretreatment years. Mid-season instantaneous gas exchange, measured on the same trees for the past four years, indicated that these responses resulted from higher needle photosynthetic rates and reduced stomatal conductance. Needles of P. sylvestris developing under increased CO₂ and temperature had consistently lower stomatal densities than their ambient grown counterparts on the same trees. The stomatal density of P. sylvestris needles was inversely correlated with δ¹³C-derived WUE, implying some effect of this morphological response on leaf gas exchange. Future atmospheric CO₂ and temperature increases are therefore likely to improve the water economy of P. sylvestris, at least at the scale of individual needles, by affecting stomatal density and gas exchange processes.     

Estimation of canopy average mesophyll conductance using δ(13) C of phloem contents

Conductance to CO(2) inside leaves, known as mesophyll conductance (g(m)), imposes large limitations on photosynthesis. Because g(m) is difficult to quantify, it is often neglected in calculations of (13)C photosynthetic discrimination. The 'soluble sugar method' estimates g(m) via differences between observed photosynthetic discrimination, calculated from the δ(13)C of soluble sugars, and discrimination when g(m) is infinite. We expand upon this approach and calculate a photosynthesis-weighted average for canopy mesophyll conductance ((c) g(m)) using δ(13)C of stem phloem contents. We measured gas exchange at three canopy positions and collected stem phloem contents in mature trees of three conifer species (Pseudotsuga menziesii, Thuja plicata and Larix occidentalis). We generated species-specific and seasonally variable estimates of (c)g(m) . We found that (c)g(m) was significantly different among species (0.41, 0.22 and 0.09 mol m(-2) s(-1) for Larix, Pseudotsuga and Thuja, respectively), but was similar throughout the season. Ignoring respiratory and photorespiratory fractionations ((c)Δ(ef)) resulted in ≈30% underestimation of (c)g(m) in Larix and Pseudotsuga, but was innocuous in Thuja. Substantial errors (~1-4‰) in photosynthetic discrimination calculations were introduced by neglecting (c)g(m) and (c)Δ(ef) . Our method is easy to apply and cost-effective, captures species variation and would have captured seasonal variation had it existed. The method provides an average canopy value, which makes it suitable for parameterization of canopy-scale models of photosynthesis, even in tall trees.     

Impacts of tree height on leaf hydraulic architecture and stomatal control in Douglas-fir

This study investigated the mechanisms involved in the regulation of stomatal closure in Douglas-fir and evaluated the potential impact of compensatory adjustments in response to increasing tree height upon these mechanisms. In the laboratory, we measured leaf hydraulic conductance (K(leaf)) as leaf water potential (Psi(l)) declined for comparison with in situ diurnal patterns of stomatal conductance (g(s)) and Psi(l) in Douglas-fir across a height gradient, allowing us to infer linkages between diurnal changes in K(leaf) and g(s). A recently developed timed rehydration technique was used in conjunction with data from pressure-volume curves to develop hydraulic vulnerability curves for needles attached to small twigs. Laboratory-measured K(leaf) declined with increasing leaf water stress and was substantially reduced at Psi(l) values of -1.34, -1.45, -1.56 and -1.92 MPa for foliage sampled at mean heights of approximately 20, 35, 44 and 55 m, respectively. In situ g(s) measurements showed that stomatal closure was initiated at Psi(l) values of -1.21, -1.36, -1.74 and -1.86 MPa along the height gradient, which was highly correlated with Psi(l) values at loss of K(leaf). Cryogenic scanning electron microscopy (SEM) images showed that relative abundances of embolized tracheids in the central vein increased with increasing leaf water stress. Leaf embolism appeared to be coupled to changes in g(s) and might perform a vital function in stomatal regulation of plant water status and water transport in conifers. The observed trends in g(s) and K(leaf) in response to changes in Psi(l) along a height gradient suggest that the foliage at the tops of tall trees is capable of maintaining stomatal conductance at more negative Psi(l). This adaptation may allow taller trees to continue to photosynthesize during periods of greater water stress.     

Tree-Ring Stable Isotopes Reveal Twentieth-Century Increases in Water-Use Efficiency of Fagus sylvatica and Nothofagus spp. in Italian and Chilean Mountains

Changes in intrinsic water use efficiency (iWUE) were investigated in Fagus sylvatica and Nothofagus spp. over the last century. We combined dendrochronological methods with dual-isotope analysis to investigate whether atmospheric changes enhanced iWUE of Fagus and Nothofagus and tree growth (basal area increment, BAI) along latitudinal gradients in Italy and Chile. Post-maturation phases of the trees presented different patterns in δ¹³C, Δ¹³C, δ¹⁸O, Ci (internal CO₂ concentration), iWUE, and BAI. A continuous enhancement in isotope-derived iWUE was observed throughout the twentieth century, which was common to all sites and related to changes in Ca (ambient CO₂ concentration) and secondarily to increases in temperature. In contrast to other studies, we observed a general increasing trend of BAI, with the exception of F. sylvatica in Aspromonte. Both iWUE and BAI were uncoupled with the estimated drought index, which is in agreement with the absence of enduring decline in tree growth. In general, δ¹³C and δ¹⁸O showed a weak relationship, suggesting the major influence of photosynthetic rate on Ci and δ¹³C, and the minor contribution of the regulation of stomatal conductance to iWUE. The substantial warming observed during the twentieth century did not result in a clear pattern of increased drought stress along these latitudinal transects, because of the variability in temporal trends of precipitation and in specific responses of populations.     

Gas exchange and hydraulic properties in the crowns of two tree species in a Panamanian moist forest

Hydraulic properties and gas exchange were measured in branches of two tropical tree species (Simarouba amara Aubl. and Tapirira guianensis Aubl.) in a moist lowland forest in Panama. Branch-level sapflow, leaf-level stomatal conductance, and water potential measurements, along with measurements of specific hydraulic conductivity of stems in crown tops, were used to relate hydraulic parameters to leaf conductance in two individuals of each species. Branches of the taller trees for each species (28 m, 31 m) showed much higher leaf-specific hydraulic conductance and leaf vapor-phase conductance than those of the smaller trees (18m, 23m). This was probably related to the leaf-to-sapwood area ratio in branches of taller trees, which was less than half that in branches of smaller trees. Dye staining showed evidence of massive cavitation in all trees, indicating that stomata do not control leaf water potential to prevent xylem cavitation in these species. Stomatal conductance of intact leaves also appeared to be insensitive to leaf area removal treatment of nearby foliage. Nevertheless, a simple mass-balance model of water flux combining hydraulic and vapor transport was in close agreement with observed maximal vapor-phase conductance in the four trees (r²=0.98, P=0.006). Our results suggest that the major organismal control over water flux in these species is by structural (leaf area) rather than physiological (stomatal) means.     

Water shortage affects the water and nitrogen balance in Central European beech forests

Whilst forest policy promotes cultivation and regeneration of beech dominated forest ecosystems, beech itself is a highly drought sensitive tree species likely to suffer from the climatic conditions prognosticated for the current century. Taking advantage of model ecosystems with cool-moist and warm-dry local climate, the latter assumed to be representative for future climatic conditions, the effects of climate and silvicultural treatment (different thinning regimes) on water status, nitrogen balance and growth parameters of adult beech trees and beech regeneration in the understorey were assessed. In addition, validation experiments with beech seedlings were carried out under controlled conditions, mainly in order to assess the effect of drought on the competitive abilities of beech. As measures of water availability xylem flow, shoot water potential, stomatal conductance as well as delta (13)C and delta (18)O in different tissues (leaves, phloem, wood) were analysed. For the assessment of nitrogen balance we determined the uptake of inorganic nitrogen by the roots as well as total N content and soluble N compounds in different tissues of adult and young trees. Retrospective and current analysis of delta (13)C, growth and meteorological parameters revealed that beech growing under warm-dry climatic conditions were impaired in growth and water balance during periods with low rain-fall. Thinning affected water, N balance and growth mostly of young beech, but in a different way under different local climatic conditions. Under cool, moist conditions, representative for the current climatic and edaphic conditions in beech forests of Central Europe, thinning improves nutrient and water status consistent to published literature and long-term experience of forest practitioners. However, beech regeneration was impaired as a result of thinning at higher temperatures and under reduced water availability, as expected in future climate.     

Twentieth century changes of tree-ring δ13C at the southern range-edge of Fagus sylvatica: increasing water-use efficiency does not avoid the growth decline induced by warming at low altitudes

We aimed to gain knowledge on the changes in intrinsic water use efficiency (iWUE) in response to increasing atmospheric CO₂ concentrations and climate change over the last century. We investigated the variation in the iWUE of mature Fagus sylvatica trees located in the higher, central and lower altitudinal forest limits (HFL, CFA and LFL) of one of the southernmost sites of beech distribution in Europe, the Montseny Mountains in Catalonia (northeast Spain), during the last century by analysing the δ¹³C of their tree rings. Pre‐ and post‐maturation phases of the trees presented different trends in δ¹³C, Δ¹³C, Ci (internal CO₂ concentration), iWUE and basal area increment (BAI). Moreover, these variables showed different trends and absolute values in the LFL than in the other altitudinal sites, CFA and HFL. Our results show the existence of an age effect on δ¹³C in the CFA and HFL (values increased by ca. 1.25‰ coinciding with the BAI suppression and release phases, previous to maturation). These age‐related changes were not found in the LFL, whose beech trees arrived to maturation earlier and experienced drier conditions during the suppression phase. In the last 26 years of comparable mature trees, the increase of iWUE deduced from the Δ¹³C analyses was ca. 10% in LFL, ca. 6% in CFA and not significant in HFL. These results show that climate change towards more arid conditions accounted for these higher Δ¹³C‐values and increases in the LFL more than the continuous increase in atmospheric CO₂ concentrations. This increased iWUE in the LFL did not avoid a decline in growth in these lowest altitudes of this beech southern range‐edge as a result of warming. Furthermore, since there was no apparent change in iWUE and growth in the beech forests growing in the more standard‐adequate environments of higher altitudes in the last 26 years, the rate of sequestration of C into temperate ecosystems may not increase with increasing atmospheric CO₂ concentrations as predicted by most models based on short‐term small scale experiments.