Comparison of whole wood and cellulose carbon and oxygen isotope series from Pinus nigra ssp. laricio (Corsica/France)

Stable isotopes in tree rings have widely been used for palaeoclimate reconstructions since tree rings record climatic information at annual resolution. However, various wood components or different parts of an annual tree-ring may differ in their isotopic compositions. Thus, sample preparation and subsequent laboratory analysis are crucial for the isotopic signal retained in the final tree-ring isotope series used for climate reconstruction and must therefore be considered for the interpretation of isotope–climate relationships. This study focuses on wood of Corsican Pine trees (Pinus nigra ssp. laricio) as this tree species allows to reconstruct the long-term climate evolution in the western Mediterranean. In a pilot study, we concentrated on methodological issues of sample preparation techniques in order to evaluate isotope records measured on pooled whole tree-ring cellulose and whole tree-ring bulk wood samples. We analysed 80-year long carbon and oxygen chronologies of Corsican Pine trees growing near the upper tree line on Corsica. Carbon and oxygen isotope records of whole tree-ring bulk wood and whole tree-ring cellulose from a pooled sample of 5 trees were correlated with the climate parameters monthly precipitation, temperature and the self-calibrating Palmer Drought Severity Index (sc-PDSI). Results show that the offsets in carbon and oxygen isotopes of bulk wood and cellulose are not constant over time. Both isotopes correlate with climate parameters from late winter and summer. The carbon and oxygen isotope ratios of cellulose are more sensitive to climatic variables than those of bulk wood. The results of this study imply that extraction of cellulose is a pre-requisite for the reconstruction of high-resolution climate records from stable isotope series of P. nigra ssp. laricio.     

Tracing carbon and oxygen isotope signals from newly assimilated sugars in the leaves to the tree-ring archive

The analysis of δ ¹³C and δ ¹⁸O in tree-ring archives offers retrospective insights into environmental conditions and ecophysiological processes. While photosynthetic carbon isotope discrimination and evaporative oxygen isotope enrichment are well understood, we lack information on how the isotope signal is altered by downstream metabolic processes.
In Pinus sylvestris , we traced the isotopic signals from their origin in the leaf water ( δ ¹⁸O) or the newly assimilated carbon ( δ ¹³C), via phloem sugars to the tree-ring, over a time-scale that ranges from hours to a growing season.
Seasonally, variable ¹³C enrichment of sugars related to phloem loading and transport did lead to uncoupling between δ ¹³C in the tree-ring, and the c _i/ c _a ratio at the leaf level. In contrast, the oxygen isotope signal was transferred from the leaf water to the tree-ring with an expected enrichment of 27‰, with time-lags of approximately 2 weeks and with a 40% exchange between organic oxygen and xylem water oxygen during cellulose synthesis.
This integrated overview of the fate of carbon and oxygen isotope signals within the model tree species P. sylvestris provides a novel physiological basis for the interpretation of δ ¹³C and δ ¹⁸O in tree-ring ecology.     

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     

Oxygen and Hydrogen Isotope Fractionation during Cellulose Metabolism in Lemna gibba L

Lemna gibba L. B3 was grown under heterotrophic, photoheterotrophic, and autotrophic conditions in water having a variety of hydrogen and oxygen isotopic compositions. The slopes of the linear regression lines between the isotopic composition of water and leaf cellulose indicated that under the three growth conditions about 40, 70, and 100% of oxygens and carbon-bound hydrogens of cellulose exchanged with those of water prior to cellulose formation. Using the equations of the linear relationships, we estimated the overall fractionation factors between water and the exchanged oxygen and carbon bound-hydrogen of cellulose. At least two very different isotope effects must determine the hydrogen isotopic composition of Lemna cellulose. One reflects the photosynthetic reduction of NADP, while the second reflects exchange reactions that occur subsequent to NADP reduction. Oxygen isotopic composition of cellulose apparently is determined by a single type of exchange reaction with water. Under different growth conditions, variations in metabolic fluxes affect the hydrogen isotopic composition of cellulose by influencing the extent to which the two isotope effects mentioned above are recorded. The oxygen isotopic composition of cellulose is not affected by such changes in growth conditions.     

Variation in oxygen isotope fractionation during cellulose synthesis: intramolecular and biosynthetic effects

The oxygen isotopic composition of plant cellulose is commonly used for the interpretations of climate, ecophysiology and dendrochronology in both modern and palaeoenvironments. Further applications of this analytical tool depends on our in-depth knowledge of the isotopic fractionations associated with the biochemical pathways leading to cellulose. Here, we test two important assumptions regarding isotopic effects resulting from the location of oxygen in the carbohydrate moiety and the biosynthetic pathway towards cellulose synthesis. We show that the oxygen isotopic fractionation of the oxygen attached to carbon 2 of the glucose moieties differs from the average fractionation of the oxygens attached to carbons 3-6 from cellulose by at least 9%, for cellulose synthesized within seedlings of two different species (Triticum aestivum L. and Ricinus communis L.). The fractionation for a given oxygen in cellulose synthesized by the Triticum seedlings, which have starch as their primary carbon source, is different than the corresponding fractionation in Ricinus seedlings, within which lipids are the primary carbon source. This observation shows that the biosynthetic pathway towards cellulose affects oxygen isotope partitioning, a fact heretofore undemonstrated. Our findings may explain the species-dependent variability in the overall oxygen isotope fractionation during cellulose synthesis, and may provide much-needed insight for palaeoclimate reconstruction using fossil cellulose.     

Divergent biochemical fractionation, not convergent temperature, explains cellulose oxygen isotope enrichment across latitudes

Recent findings based on the oxygen isotope ratios of tree trunk cellulose indicate that the temperature of biomass production in biomes ranging from boreal to subtropical forests converge to an average leaf temperature of 21.4°C. The above conclusion has been drawn under the assumption that biochemically related isotopic fractionations during cellulose synthesis are not affected by temperature. Here we test the above assumption by heterotrophically generating cellulose at different temperatures and measuring the proportion of carbohydrate oxygen that exchange with water during cellulose synthesis and the average biochemical fractionation associated with this exchange. We observed no variation in the proportion of oxygen that exchange with different temperatures, which averaged 0.42 as it has been observed in other studies. On the other hand, the biochemical oxygen isotope fractionation during cellulose synthesis is affected by temperature and can be described by a 2(nd) order polynomial equation. The biochemical fractionation changes little between temperatures of 20 and 30°C averaging 26‰ but increases at lower temperatures to values of 31‰. This temperature sensitive biochemical fractionation explains the pattern of cellulose oxygen isotope ratios of aquatic plants encompassing several latitudes. The observed temperature sensitive biochemical fractionation also indicates that divergent biochemical fractionation and not convergent leaf temperature explains the increase in oxygen isotope enrichment of cellulose across several biomes.     

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.     

Identifying drivers of leaf water and cellulose stable isotope enrichment in <Emphasis Type="Italic">Eucalyptus</Emphasis> in northern Australia

Several previous studies have investigated the use of the stable hydrogen and oxygen isotope compositions in plant materials as indicators of palaeoclimate. However, accurate interpretation relies on a detailed understanding of both physiological and environmental drivers of the variations in isotopic enrichments that occur in leaf water and associated organic compounds. To progress this aim we measured δ¹⁸O and δ²H values in eucalypt leaf and stem water and δ¹⁸O values in leaf cellulose, along with the isotopic compositions of water vapour, across a north-eastern Australian aridity gradient. Here we compare observed leaf water enrichment, along with previously published enrichment data from a similar north Australian transect, to Craig–Gordon-modelled predictions of leaf water isotopic enrichment. Our investigation of model parameters shows that observed ¹⁸O enrichment across the aridity gradients is dominated by the relationship between atmospheric and internal leaf water vapour pressure while ²H enrichment is driven mainly by variation in the water vapour—source water isotopic disequilibrium. During exceptionally dry and hot conditions (RH < 21%, T > 37 °C) we observed strong deviations from Craig–Gordon predicted isotope enrichments caused by partial stomatal closure. The atmospheric–leaf vapour pressure relationship is also a strong predictor of the observed leaf cellulose δ¹⁸O values across one aridity gradient. Our finding supports a wider applicability of leaf cellulose δ¹⁸O composition as a climate proxy for atmospheric humidity conditions during the leaf growing season than previously documented.     

Climatic significance of isotope ratios

The hydrogen and oxygen isotope ratios of water, which can be measured by Isotope Ratio Mass Spectrometry (IRMS), exhibit climatic dependencies and are commonly exploited in hydrogeology. More generally, the overall carbon or hydrogen isotope ratios of plant organic matter, and in particular of tree-ring cellulose, have been frequently used for climatic reconstruction. However, since many physicochemical and biochemical fractionation phenomena are likely to contribute to the isotopic values, the interpretation of the climatic significance of isotopic parameters is not always straightforward. In the case of hydrogen and oxygen for instance, the climatic profile of the source meteoric water is not simply transferred to leaf water and many steps of the biosyntheses are accompanied by kinetic and thermodynamic isotope effects that depend on the individual mechanistic pathways. The information brought about by overall isotope ratios determined by IRMS is averaged over all fractionation effects undergone at the different molecular positions. In contrast, the NMR investigation of Site-specific Natural Isotope Fractionation (SNIF-NMR) gives simultaneous access to isotope ratios specific to individual positions in the molecule. Since the different atoms do not necessarily exhibit the same climatic dependency, the method provides complementary responses to the environmental conditions. In particular, the isotopic parameters of ethanol and water obtained by fermenting sugars in standardized conditions reflect climatic influences which took place at different periods of plant growth. As a consequence, statistical analyses based on multi-site isotopic variables provide powerful criteria for distinguishing geographical regions of cultivation characterized by different climatic features. Although the sensitivity to climatic variations is the most pronounced for plant water and for sugars formed at the first step of photosynthesis, other components such as lipids or minor metabolites also exhibit climatic dependencies. The combination of isotopic values pertaining to different atomic species and either averaged over the whole molecule (IRMS) or associated with different molecular sites (SNIF-NMR), provides complementary criteria, which can be exploited in terms of both climatic significance and mechanistic pathways of the individual atoms.     

Tree-ring widths and isotopes of artificially defoliated balsam firs: A simulation of spruce budworm outbreaks in Eastern Canada

Defoliation by insects is a major disturbance influencing the forest dynamics in many ecosystems and can affect forest productivity worldwide. The main objective of this research was to further investigate the potential use of tree-ring widths and isotopic compositions to identify different degrees of past spruce budworm defoliation episodes. A secondary objective was to understand the responses of trees to defoliation episodes using carbon isotopes as a proxy to provide insights into subsequent physiological changes. Tree-ring widths, carbon and oxygen isotopic compositions in wood cellulose and gas exchange measurements were compared among 288 balsam fir (Abies balsamea Mill.) seedlings grown in a controlled experiment that involved different intensities of defoliation. Observations were performed over four growing periods. Moderate to heavy-defoliated seedlings showed reduced radial growth and enriched their cellulose carbon isotopic composition probably as a result of mobilized stored carbohydrates enriched in ¹³C. Less severely defoliated seedlings did not show significant reductions in growth and ¹³C enrichments. The gas exchange observations and wood cellulose oxygen isotope compositions do not suggest photosynthetic compensation in the remaining needles although a positive trend in the response of both assimilation rate (A) and stomatal conductance (g_s) to defoliation was observed in the first growing period. Thus it remains open as to which mechanisms were employed to compensate for the reduced carbon source in the mildly defoliated seedlings. While further investigations are advised, the results of this study still help promote the utilization of tree-ring widths in combination with carbon isotopic compositions for reconstructing severe past defoliation events.     

A simplified GIS approach to modeling global leaf water isoscapes

The stable hydrogen (delta(2)H) and oxygen (delta(18)O) isotope ratios of organic and inorganic materials record biological and physical processes through the effects of substrate isotopic composition and fractionations that occur as reactions proceed. At large scales, these processes can exhibit spatial predictability because of the effects of coherent climatic patterns over the Earth's surface. Attempts to model spatial variation in the stable isotope ratios of water have been made for decades. Leaf water has a particular importance for some applications, including plant organic materials that record spatial and temporal climate variability and that may be a source of food for migrating animals. It is also an important source of the variability in the isotopic composition of atmospheric gases. Although efforts to model global-scale leaf water isotope ratio spatial variation have been made (especially of delta(18)O), significant uncertainty remains in models and their execution across spatial domains. We introduce here a Geographic Information System (GIS) approach to the generation of global, spatially-explicit isotope landscapes (= isoscapes) of "climate normal" leaf water isotope ratios. We evaluate the approach and the resulting products by comparison with simulation model outputs and point measurements, where obtainable, over the Earth's surface. The isoscapes were generated using biophysical models of isotope fractionation and spatially continuous precipitation isotope and climate layers as input model drivers. Leaf water delta(18)O isoscapes produced here generally agreed with latitudinal averages from GCM/biophysical model products, as well as mean values from point measurements. These results show global-scale spatial coherence in leaf water isotope ratios, similar to that observed for precipitation and validate the GIS approach to modeling leaf water isotopes. These results demonstrate that relatively simple models of leaf water enrichment combined with spatially continuous precipitation isotope ratio and climate data layers yield accurate global leaf water estimates applicable to important questions in ecology and atmospheric science.     

Summer temperature dependency of larch budmoth outbreaks revealed by Alpine tree-ring isotope chronologies

Larch budmoth (LBM, Zeiraphera diniana Gn.) outbreaks cause discernable physical alteration of cell growth in tree rings of host subalpine larch (Larix decidua Mill.) in the European Alps. However, it is not clear if these outbreaks also impact isotopic signatures in tree-ring cellulose, thereby masking climatic signals. We compared LBM outbreak events in stable carbon and oxygen isotope chronologies of larch and their corresponding tree-ring widths from two high-elevation sites (1800–2200 m a.s.l.) in the Swiss Alps for the period AD 1900–2004 against isotope data obtained from non-host spruce (Picea abies). At each site, two age classes of tree individuals (150–250 and 450–550 years old) were sampled. Inclusion of the latter age class enabled one chronology to be extended back to AD 1650, and a comparison with long-term monthly resolved temperature data. Within the constraints of this local study, we found that: (1) isotopic ratios in tree rings of larch provide a strong and consistent climatic signal of temperature; (2) at all sites the isotope signatures were not disturbed by LBM outbreaks, as shown, for example, by exceptionally high significant correlations between non-host spruce and host larch chronologies; (3) below-average July to August temperatures and LBM defoliation events have been coupled for more than three centuries. Dampening of Alps-wide LBM cyclicity since the 1980s and the coincidence of recently absent cool summers in the European Alps reinforce the assumption of a strong coherence between summer temperatures and LBM defoliation events. Our results demonstrate that stable isotopes in tree-ring cellulose of larch are an excellent climate proxy enabling the analysis of climate-driven changes of LBM cycles in the long term.     

Effects of environmental parameters, leaf physiological properties and leaf water relations on leaf water delta18O enrichment in different Eucalyptus species

Stable oxygen isotope ratios (delta18O) have become a valuable tool in the plant and ecosystem sciences. The interpretation of delta18O values in plant material is, however, still complicated owing to the complex interactions among factors that influence leaf water enrichment. This study investigated the interplay among environmental parameters, leaf physiological properties and leaf water relations as drivers of the isotopic enrichment of leaf water across 17 Eucalyptus species growing in a common garden. We observed large differences in maximum daily leaf water delta18O across the 17 species. By fitting different leaf water models to these empirical data, we determined that differences in leaf water delta18O across species are largely explained by variation in the Péclet effect across species. Our analyses also revealed that species-specific differences in transpiration do not explain the observed differences in delta18O while the unconstrained fitting parameter 'effective path length' (L) was highly correlated with delta18O. None of the leaf morphological or leaf water related parameters we quantified in this study correlated with the L values we determined even though L was typically interpreted as a leaf morphological/anatomical property. A sensitivity analysis supported the importance of L for explaining the variability in leaf water delta18O across different species. Our investigation highlighted the importance of future studies to quantify the leaf properties that influence L. Obtaining such information will significantly improve our understanding of what ultimately determines the delta18O values of leaf water across different plant species.     

Comparison of measured and modeled variations in piñon pine leaf water isotopic enrichment across a summer moisture gradient

Stable hydrogen and oxygen isotopic composition of bulk leaf water (δD_lw and δ¹⁸O_lw) in piñon pine (Pinus edulis and P. monophylla) and gas exchange parameters were measured under field conditions to examine the effects of seasonal moisture stress on leaf water isotopic enrichment. Study sites were located near the lower elevation limit for piñon in the southwestern USA. Leaf-level transpiration measurements were made four times daily in spring, summer and early autumn; simultaneously, leaf samples were collected for water extraction and stable isotope analysis. Diurnal variations in δD_lw and δ¹⁸O_lw values were small, especially when leaf water residence times (molar leaf water content divided by transpiration rate) were high. Stomatal conductance explained most of the variance (60%) in leaf water enrichment across the dataset. Observed leaf water enrichment was compared with predictions of steady-state and nonsteady-state models. Nonsteady-state predictions fit observations the best, although D enrichment was often lower than predicted by any model. Hydrogen isotope ratios of leaf water and cellulose nitrate were strongly correlated, demonstrating preservation of a leaf water signal in wood and leaf cellulose.     

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     

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

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

Variations in the natural abundance of oxygen-18 and deuterium in plant carbohydrates

Variations in the natural abundance of ¹⁸O and ²H in plant cellulose are influenced by the isotopic composition of the water directly involved in metabolism—the metabolic water fraction. The isotopic distinction between the metabolic source water and total tissue water must reflect the formation of isotopic gradients within the tissue that are influenced by the rate of water turnover, by properties of the water conducting system and by environmental conditions. It seems that the ¹⁸O abundance in the metabolic water is conserved in cellulose with a relatively constant isotope effect. The relationship of the ²H abundance between metabolic water and cellulose is more complex. Hydrogen incorporated into photosynthetic products during primary reduction steps is highly depleted in ²H. However, a large proportion of these hydrogens are subsequently replaced by exchange with water, leading to ²H enrichment during heterotrophic metabolism. Deciphering the oxygen isotope ratio of cellulose could help in providing insights into the carbon and oxygen fluxes exchanged between plants and the atmosphere. This is because the ¹⁸O abundance in cellulose records the ¹⁸O abundance in the metabolic water, which in turn, controls the oxygen isotopic signatures of the CO₂ and O₂ released by plants into the atmosphere. The hydrogen isotope effects associated with carbohydrate metabolism provide insights into the autotrophic state of a plant tissue. This is because the hydrogen isotope ratio of carbohydrates must reflect the net effects of the two opposing isotope effects associated with photosynthesis and heterotrophic metabolism.     

What can we Learn from Oxygen and Hydrogen Isotope Compositions of Tree-ring Cellulose in Higher Plants (英文)

Stable isotope ratios in tree-ring cellulose have been shown to be reliable recorders of changes in the ambient climate (Leuenberger et al., 1998). Thus, isotopic fractionations associated with both physical and biochemical processes during cellulose synthesis in higher plants (Epstein & Krishnaumurthy, 1990; Roden et al., 2000; Saurer et al., 1997a; 1997b; 2000) can be used as archives for past climatic indicators. Superimposed on the climatic induced isotopic signal are the long-term responses of plant physiological processes to past changes in environmental conditions including CO2 enrichment.     

Effects of Mild Water Stress and Diurnal Changes in Temperature and Humidity on the Stable Oxygen and Hydrogen Isotopic Composition of Leaf Water in Cornus stolonifera L

In this paper we make comparisons between the observed stable isotopic composition of leaf water and the predictions of the Craig-Gordon model of isotopic enrichment when plants (Cornus stolonifera L.) were exposed to natural, diurnal changes in temperature and humidity in a glasshouse. In addition, we determined the effects of mild water stress on the isotopic composition of leaf water. The model predicted different patterns of diurnal change for the oxygen and hydrogen isotopic composition of leaf water. The observed leaf water isotopic composition followed qualitatively similar patterns of diurnal change to those predicted by the model. At midday, however, the model always predicted a higher degree of heavy isotope enrichment than was actually observed in leaves. There was no effect of mild water stress on the hydrogen isotopic composition of leaf water. For the oxygen isotopic composition of leaf water, there was either no significant difference between control and water-stressed plants or the stressed plants had lower δ¹⁸O values, despite the enriched stem water isotopic composition observed for the stressed plants.     

Consequences of larch budmoth outbreaks on the climatic significance of ring width and stable isotopes of larch

Tree-ring widths and stable carbon and oxygen isotopes of five European larch trees from Lötschental, Switzerland were investigated for the period 1900–2004. The objective was to test the suitability of each of these parameters for high-frequency climate reconstructions. This is of special interest with regard to the problem of cyclic larch budmoth (LBM) infestations of alpine larch trees. The results clearly demonstrate that tree-ring width chronologies are not suitable for high-frequency reconstructions because infestations lead to variably reduced tree-ring increments, largely suppressing climate signals. On the other hand, the stable isotope chronologies proved less affected by larch budmoth outbreaks, independent of the strength of the infestations. The correlation of the carbon isotopes with summer temperatures was especially high (r = 0.73) and with precipitation lower but nevertheless significant (r = ?0.43). Oxygen isotopes were also correlated with summer temperature (r = 0.46); however, a certain perturbation of normal oxygen isotope signatures due to LBM outbreaks was evident. Contrary to tree-ring widths, none of the LBM outbreaks caused a significant disturbance of the current year’s isotopic climate signal and, most importantly, there were no delayed effects in the following years. Thus, stable carbon isotopes in tree-ring chronologies of the European larch provide an excellent opportunity for high-frequency temperature reconstructions.