Aerial decay influence on the stable oxygen and carbon isotope ratios in tree ring cellulose

Sub-fossil wood is often affected by the decaying process that introduces uncertainties in the measurement of oxygen and carbon stable isotope composition in cellulose. Although the cellulose stable isotopes are widely used as climatic proxies, our understanding of processes controlling their behavior is very limited. We present here a comparative study of stable oxygen and carbon isotope ratios in tree ring cellulose in decayed and non-decayed wood samples of Swiss stone pine (Pinus cembra) trees. The intra-ring stable isotope variability (around the circumference of a single ring) was between 0.1 and 0.5‰ for δ¹⁸O values and between 0.5 and 1.6‰ for δ¹³C values for both decayed and non-decayed wood. Observed intra-tree δ¹⁸O variability is less than that reported in the literature (0.5–1.5‰), however, for δ¹³C it is larger than the reported values (0.7–1.2‰). The inter-tree variability for non-decayed wood ranges between 1.1 and 2.3‰ for δ¹⁸O values, and between 2 and 4.7‰ for δ¹³C values. The inter-tree differences for δ¹⁸O values are similar to those reported in the literature (1–2‰ for oxygen and 1–3‰ for carbon) but are larger for δ¹³C values. We have found that the differences for δ¹⁸O and δ¹³C values between decayed and non-decayed wood are smaller than the variation among different trees from the same site, suggesting that the decayed wood can be used for isotopic paleoclimate research.     

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.     

Do centennial tree-ring and stable isotope trends of Larix gmelinii (Rupr.) Rupr. indicate increasing water shortage in the Siberian north?

Tree-ring width of Larix gmelinii (Rupr.) Rupr., ratios of stable isotopes of C (δ¹³C) and O (δ¹⁸O) of whole wood and cellulose chronologies were obtained for the northern part of central Siberia (Tura, Russia) for the period 1864–2006. A strong decrease in the isotope ratios of O and C (after atmospheric δ¹³C corrections) and tree-ring width was observed for the period 1967–2005, while weather station data show a decrease in July precipitation, along with increasing July air temperature and vapor pressure deficit (VPD). Temperature at the end of May and the whole month of June mainly determines tree radial growth and marks the beginning of the vegetation period in this region. A positive correlation between tree-ring width and July precipitation was found for the calibration period 1929–2005. Positive significant correlations between C isotope chronologies and temperatures of June and July were found for whole wood and cellulose and negative relationships with July precipitation. These relationships are strengthened when the likely physiological response of trees to increased CO₂ is taken into account (by applying a recently developed δ¹³C correction). For the O isotope ratios, positive relationships with annual temperature, VPD of July and a negative correlation with annual precipitation were observed. The δ¹⁸O in tree rings may reflect annual rather than summer temperatures, due to the late melting of the winter snow and its contribution to the tree water supply in summer. We observed a clear change in the isotope and climate trends after the 1960s, resulting in a drastic change in the relationship between C and O isotope ratios from a negative to a positive correlation. According to isotope fractionation models, this indicates reduced stomatal conductance at a relatively constant photosynthetic rate, as a response of trees to water deficit for the last half century in this permafrost region.     

Effect of climatic variability on δ13C and tree-ring growth in piñon pine (Pinus edulis)

Understanding the response of long-lived species to natural climatic variability at multiple scales is a prerequisite for forecasting ecosystem responses to global climate change. This study investigated the response of piñon pine (Pinus edulis) to natural climatic variability using information on physiology and growth as recorded in leaves and tree rings. δ¹³C of annual leaf cohorts (δ¹³C_leaf) and tree rings (δ¹³C_ring) were measured at an ecotonal/xeric site and a mid-range/mesic site. Ring width indices (RWI) were used to estimate annual growth of individual trees. Relationships between seasonal and annual climate parameters and δ¹³C and growth were investigated. δ¹³C–climate relationships were stronger for δ¹³C_leaf than for δ¹³C_ring especially at the xeric site. The mean monthly maximum summer temperatures over May through September (summer T _max) had the strongest influence on δ¹³C_leaf. There was a strong negative relationship between RWI with summer T _max and a strong positive relationship between RWI with October to October precipitation (water–year PPN) at both sites. This suggests that piñon pine populations could be vulnerable to decreased growth and, perhaps mortality, in response to warmer, drier conditions predicted by models of global climate change.     

Tree-ring wood anatomy and stable isotopes show structural and functional adjustments in olive trees under different water availability

Background and Aims

Olive tree (Olea europaea L.) is a drought-tolerant tree species cultivated in Mediterranean-type environments. Although it is tolerant to drought, dry conditions decrease its productivity. A thorough analysis of the hydraulic architecture and wood anatomical plasticity, as well as of their physiological significance, is needed to understand how olive trees will adapt to the predicted increase in frequency and severity of drought in the Mediterranean region.

Methods

Dendrochronological, stable isotopic (δ¹³C, δ¹⁸O) and wood anatomical analyses were applied to understand how different water availability can affect wood stem structure and function, in rainfed and irrigated at 100 % of crop evapotranspiration (ETc) olive trees in an experimental orchard close to Benevento (Italy) from 1992 to 2009.

Results

Dendrochronological data indicate that cross-dating and synchronization of ring-width time series in olive tree is possible. After the start of irrigation, significantly more negative δ¹³C and lower δ¹⁸O values were recorded in irrigated trees indicating higher stomatal conductance and transpiration rates. Increased water balance induced the formation of a higher number of vessels with higher diameter.

Conclusions

Water balance variations affected wood anatomy and isotopic composition. Anatomical analyses detected structural and functional adjustments in rainfed trees that produced more vessels with lower diameter to prevent cavitation. Isotopic analyses confirmed that irrigated trees continuously showed enhanced transpiration rates.     

Tracing plant source water dynamics during drought by continuous transpiration measurements: An in-situ stable isotope approach

The isotopic composition of xylem water (δ_X) is of considerable interest for plant source water studies. In-situ monitored isotopic composition of transpired water (δ_T) could provide a nondestructive proxy for δ_X-values. Using flow-through leaf chambers, we monitored 2-hourly δ_T-dynamics in two tropical plant species, one canopy-forming tree and one understory herbaceous species. In an enclosed rainforest (Biosphere 2), we observed δ_T-dynamics in response to an experimental severe drought, followed by a ²H deep-water pulse applied belowground before starting regular rain. We also sampled branches to obtain δ_X-values from cryogenic vacuum extraction (CVE). Daily flux-weighted δ¹⁸O_T-values were a good proxy for δ¹⁸O_X-values under well-watered and drought conditions that matched the rainforest's water source. Transpiration-derived δ¹⁸O_X-values were mostly lower than CVE-derived values. Transpiration-derived δ²H_X-values were relatively high compared to source water and consistently higher than CVE-derived values during drought. Tracing the ²H deep-water pulse in real-time showed distinct water uptake and transport responses: a fast and strong contribution of deep water to canopy tree transpiration contrasting with a slow and limited contribution to understory species transpiration. Thus, the in-situ transpiration method is a promising tool to capture rapid dynamics in plant water uptake and use by both woody and nonwoody species.     

Trophic position and seasonal changes in the diet of the red wood ant Formica aquilonia as indicated by stable isotope analysis

1. Red wood ants are among the most numerous generalist predators and strongly affect the composition of arthropod communities in forest ecosystems. However, their trophic position remains poorly understood. Stable isotope analysis was applied to study the trophic position of Formica aquilonia and reveal seasonal changes in its trophic links with both myrmecophilous aphids and other invertebrates in a mixed forest of western Siberia. 2. The δ¹⁵N values of F. aquilonia exceeded those of herbivores and aphids by approximately 3.5‰. Despite obligate trophobiotic relationships with aphids, F. aquilonia occupied the trophic position of first‐order predator. The higher content of ¹³C in the worker ants, compared with members of grazing food chains, was explained by their consumption of ¹³C‐enriched aphid honeydew. 3. Myrmecophilous tree‐dwelling aphids were enriched in ¹³C and ¹⁵N relative to grass‐inhabiting species, and the honeydew of tree‐dwelling aphids had higher δ¹³C values than those of the honeydew of grass‐inhabiting aphids. 4. The decrease in δ¹³C values of the worker ants from spring and summer to autumn apparently reflected the transition from the collection of tree sap and feeding on the aphid honeydew from trees with high ¹³C content in the spring and early summer to a more diverse liquid diet in late summer, which included ¹³C‐depleted honeydew of aphids from herbs. 5. The prevalence of the ¹⁵N‐depleted aphid honeydew in the ants' diet in the second half of the summer is discussed as one possible explanation for the seasonal decline in δ¹⁵N values of the worker ants.     

Carbon Isotope Discrimination is not Correlated with Transpiration Efficiency in Three Cool-Season Grain Legumes （Pulses）

The carbon isotope discrimination （δ^13C） of leaves has been shown to be correlated with the transpiration efficiency of leaves in a wide range of species. This has led to δ^13C being used in breeding programs to select for improved transpiration efficiency. The correlation between δ^13C and transpiration efficiency was determined under well-watered conditions during the vegetative phase in six genotypes of lentil （Lens culinaris Medikus）, six genotypes of chickpea （Cicerarietinum L.） and 10 cultivars of narrow-leafed lupin （Lupinus angustifolius L.）. Biomass （dry matter） accumulation and water use （transpiration） varied among the genotypes in all three species and transpiration efficiency was 40% to 75% higher in the most efficient compared with the least efficient genotypes. However, δ^13C and transpiration efficiency were not significantly correlated in any of the species. This suggests that the δ^13C technique cannot be used in selection for transpiration efficiency in the three grain legumes （pulses） studied.     

Carbon, nitrogen, and hydrogen isotope ratios in creekside trees in western Kansas

Three species of creekside trees were monitored weekly during the 2007 and 2008 growing seasons. The 2007 growing season was wet early, but became drier as the season progressed. In contrast, the 2008 growing season was dry early, but became wetter as the season progressed. Creekside trees were measured to determine effects of changing water regimes on leaf-level processes. Lonicera tatarica plants were compared to Morus alba and Celtis occidentalis trees. Leaves were monitored for changes in stomatal conductance, transpiration, δ¹³C, δ¹⁵N, δD, leaf temperature, and heat losses via latent, sensible, and radiative pathways. δD of creek water was more similar to ground water than to rain water, but the creek was partially influenced by summer rains. δD of bulk leaf material was significantly higher in individuals of C. occidentalis compared to the other species, consistent with source water coming from shallower soil layers. Despite decreasing water levels, none of these tree species showed signs of water stress. There were no significant differences between species in stomatal conductance or transpiration. Leaf δ¹³C was significantly lower in individuals of L. tatarica compared to the other species. Differences in δ¹³C were attributed to a lower carboxylation capacity, consistent with lower leaf nitrogen content in L. tatarica plants. Leaf δ¹⁵N was significantly lower in individuals of L. tatarica compared to the other species, consistent with uptake of a different N source. Two of the three sites appeared to be affected by inorganic N from fertilizer run-off. Evidence is presented that these species acquired water and nitrogen from different sources, resulting from differences in root uptake patterns.     

Carbon isotope compositions (δ13C) of leaf,wood and holocellulose differ among genotypes of poplar and between previous land uses in a short‐rotation biomass plantation

The efficiency of water use to produce biomass is a key trait in designing sustainable bioenergy‐devoted systems. We characterized variations in the carbon isotope composition (δ¹³C) of leaves, current year wood and holocellulose (as proxies for water use efficiency, WUE) among six poplar genotypes in a short‐rotation plantation. Values of δ¹³C_wood and δ¹³C_{holocellulose} were tightly and positively correlated, but the offset varied significantly among genotypes (0.79–1.01‰). Leaf phenology was strongly correlated with δ¹³C, and genotypes with a longer growing season showed a higher WUE. In contrast, traits related to growth and carbon uptake were poorly linked to δ¹³C. Trees growing on former pasture with higher N‐availability displayed higher δ¹³C as compared with trees growing on former cropland. The positive relationships between δ¹³C_leaf and leaf N suggested that spatial variations in WUE over the plantation were mainly driven by an N‐related effect on photosynthetic capacities. The very coherent genotype ranking obtained with δ¹³C in the different tree compartments has some practical outreach. Because WUE remains largely uncoupled from growth in poplar plantations, there is potential to identify genotypes with satisfactory growth and higher WUE.     

Climate at the onset of western Mediterranean agriculture expansion: Evidence from stable isotopes of sub-fossil oak tree rings in Spain

Climate conditions during the early Holocene may have contributed to the spread of Old World agriculture from its area of birth, in the Fertile Crescent (Near East), west through the Mediterranean Basin. Reconstructing the specific environmental conditions existing in early agricultural settlements of the western Mediterranean may help to elucidate this point. The aim of this work was to gain information on past climate of one of the earliest agricultural settlements of the Iberian Peninsula, La Draga, a lacustrine site dating back to the 2nd half of the 8th millennium BP, in which post fragments of deciduous oak have been exceptionally well preserved in an anaerobic environment (sub-fossil wood). We studied the relationship between climate factors and carbon isotope discrimination (Δ¹³C) and oxygen isotope composition (δ¹⁸O) in wood α-cellulose from modern tree-ring series of Quercus humilis Mill. and Quercus petraea (Matt) Liebl. recovered from the area. Climatic responses observed in extant material were used to interpret the isotopic signal of wood α-cellulose extracted from sub-fossil wood. Results showed that water availability of late spring, early summer and September influences Δ¹³C and δ¹⁸O, which allows their use as proxies for palaeoclimatic reconstruction. Differences between sub-fossil and extant samples in Δ¹³C (19.35 vs. 18.02‰) and δ¹⁸O (26.32 vs. 29.28‰) records suggest slightly lower temperatures and higher plant water availability than at present during the establishment of agriculture at the site. These results seem to disprove the existence of an environmental limitation preventing continuation of hunting and gathering activities as a cause for the adoption of agriculture in this early agricultural site on the Iberian Peninsula.     

Leaf stable carbon isotope composition reflects transpiration efficiency in Zea mays

The increasing demand for food production and predicted climate change scenarios highlight the need for improvements in crop sustainability. The efficient use of water will become increasingly important for rain‐fed agricultural crops even in fertile regions that have historically received ample precipitation. Improvements in water‐use efficiency in Zea mays have been limited, and warrant a renewed effort aided by molecular breeding approaches. Progress has been constrained by the difficulty of measuring water‐use in a field environment. The stable carbon isotope composition (δ¹³C) of the leaf has been proposed as an integrated signature of carbon fixation with a link to stomatal conductance. However, additional factors affecting leaf δ¹³C exist, and a limited number of studies have explored this trait in Z. mays. Here we present an extensive characterization of leaf δ¹³C in Z. mays. Significant variation in leaf δ¹³C exists across diverse lines of Z. mays, which we show to be heritable across several environments. Furthermore, we examine temporal and spatial variation in leaf δ¹³C to determine the optimum sampling time to maximize the use of leaf δ¹³C as a trait. Finally, our results demonstrate the relationship between transpiration and leaf δ¹³C in the field and the greenhouse. Decreasing transpiration and soil moisture are associated with decreasing leaf δ¹³C. Taken together these results outline a strategy for using leaf δ¹³C and reveal its usefulness as a measure of transpiration efficiency under well‐watered conditions rather than a predictor of performance under drought.     

Long-term transpiration in two eucalypt species in a native woodland estimated by the heat-pulse technique

Abstract The heat-pulse method was used to estimate transpiration rates continuously for periods up to 2 years in mature trees of Eucalyptus wandoo and Eucalyptus salmonophloia at two topographic locations in a remnant native woodland in the Western Australian wheatbelt. Annual transpiration per tree ranged from about 11400 to 18000 L per tree. Highest transpiration rates occurred in late spring or early summer, depending on rainfall distribution. The trees were able to rapidly utilize water following heavy rain outside the agricultural growing season. Extrapolating transpiration rates from single trees to an area of woodland showed that annual transpiration at the ridge site was 150 mm and 168 mm at a site alongside a drainage line. Scaling up transpiration from individual trees requires caution and should allow for variability in trees and soils. The role of trees in curtailing salinization is discussed.     

Influence of precipitation seasonality on piñon pine cellulose δD values

The influence of seasonal to interannual climate variations on cellulose hydrogen isotopic composition (δD) was assessed by analysing tree rings and needles of piñon pine (Pinus edulis and P. monophylla). Sites spanned a gradient of decreasing summer precipitation, from New Mexico to Arizona to Nevada. Tree rings were divided into earlywood, latewood and whole‐year increments, and annual cohorts of needles were collected. The study period (1989–96) included two La Niña events (1989, 1996) and a prolonged El Niño event (1991–95). Winter and spring moisture conditions were strongly related to October–March Southern Oscillation Index (SOI) in New Mexico and Arizona, with above‐average precipitation occurring in El Niño years. Wood δD values at these sites were correlated with winter and spring moisture conditions. Needle δD values were correlated with summer moisture conditions in New Mexico and with winter moisture and SOI in Arizona. Low cellulose δD values observed from 1991 to 1993 in both wood and needles occurred during wet El Niño years, whereas high δD values in needles were present during the dry, La Niña years of 1989 and 1996. North‐eastern Nevada does not receive precipitation anomalies related to ENSO, and thus cellulose δD values did not reflect the ENSO pattern observed at the other sites. Cellulose δD values were strongly, inversely correlated with relative humidity variations at all sites, as predicted by a mechanistic model. Contrary to predictions from the same model and observations from more mesic areas, time series of cellulose δD values were not directly correlated with interannual or seasonal variations in precipitation δD values or temperature at any of the sites. On a regional basis, however, mean δD values in needles and wood were correlated with mean annual temperature and δD values of precipitation. This suggests that temporal averaging may bias relationships between biological systems and climate.     

Rapid recycling of triose phosphates in oak stem tissue

We report the carbon-13 and oxygen-18 isotope ratios in cellulose from the early and late wood of pedunculate oak (Quercus robur L.). The δ¹³ C value of the early wood correlates best with that of the late wood of the previous year. The δ¹⁸O value of the early wood correlates best with that of the late wood of the same year. We suggest that a biochemical explanation of these data is that there is a rapid cycle between hexose monophosphates and triose phosphates in oak stem tissue during cellulose synthesis. Evidence in support of this explanation is provided by the intramolecular distribution of ¹⁴C in labelled fructose extracted from cores of wood that had been supplied with [1?¹⁴C]- and [6-¹⁴C]glucose.     

Changes in leaf photosynthetic characteristics and water use efficiency along with tree height of 4 tree species

As main photosynthetic organs, leaves are very sensitive to exterior environments. Water deficiency obviously affects the biological and physiological characteristics of leaves. Xylem pathways increase when trees grow tall, which results in the increase in water gravity as well as pathway resistance. Accordingly, the physiological characteristics of leaves change along with tree height. In this research, the photosynthetic characteristics and carbon isotope ratio (δ¹³C) in the leaves of 4 tree species, Platanus hispanica, Robinia pseudoacacia, Fraxinus chinensis and Ginkgo biloba, were measured. The results showed that the leaf photosynthetic rate (Pn), transpiration rate (Tr), stomatal conductance (Cond) and internal CO₂ concentration (Ci) reduced along with tree height, while the leaf δ¹³C increased along with tree height. The One Way ANOVA and LSD tests showed that the leaf photosynthetic characteristics and δ¹³C varied significantly at different tree heights (P < 0.05). The decrease in leaf photosynthetic capability and the increase in δ¹³C along with tree height indicate that the leaves at the tree tops suffer from water stress. These results support the hydraulic limitation hypothesis.     

Modelling non‐steady‐state isotope enrichment of leaf water in a gas‐exchange cuvette environment

The combined use of a gas‐exchange system and laser‐based isotope measurement is a tool of growing interest in plant ecophysiological studies, owing to its relevance for assessing isotopic variability in leaf water and/or transpiration under non‐steady‐state (NSS) conditions. However, the current Farquhar & Cernusak (F&C) NSS leaf water model, originally developed for open‐field scenarios, is unsuited for use in a gas‐exchange cuvette environment where isotope composition of water vapour (δ_v) is intrinsically linked to that of transpiration (δ_E). Here, we modified the F&C model to make it directly compatible with the δ_v–δ_E dynamic characteristic of a typical cuvette setting. The resultant new model suggests a role of ‘net‐flux’ (rather than ‘gross‐flux’ as suggested by the original F&C model)‐based leaf water turnover rate in controlling the time constant (τ) for the approach to steady sate. The validity of the new model was subsequently confirmed in a cuvette experiment involving cotton leaves, for which we demonstrated close agreement between τ values predicted from the model and those measured from NSS variations in isotope enrichment of transpiration. Hence, we recommend that our new model be incorporated into future isotope studies involving a cuvette condition where the transpiration flux directly influences δ_v. There is an increasing popularity among plant ecophysiologists to use a gas‐exchange system coupled to laser‐based isotope measurement for investigating non‐steady state (NSS) isotopic variability in leaf water (and/or transpiration); however, the current Farquhar & Cernusak (F&C) NSS leaf water model is unsuited for use in a gas‐exchange cuvette environment due to its implicit assumption of isotope composition of water vapor (δ_v) being constant and independent of that of transpiration (δ_E). In the present study, we modified the F&C model to make it compatible with the dynamic relationship between δ_v and δ_E as is typically associated with a cuvette setting. Using an experiment conducted on cotton leaves, we show that the modified NSS model performed well in predicting the time constant for the exponential approach of leaf water toward steady state under cuvette conditions. Such a result demonstrates the applicability of this new model to gas‐exchange cuvette conditions where the transpiration flux directly influences δ_v, and therefore suggests the need to incorporate this model into future isotope studies that employ a laser‐cuvette coupled system.     

Exploring climate forcing of growth depression in subfossil South Swedish bog pines using stable isotopes

Comparison between growth variability, based on ring-width (RW) analysis, and moisture-sensitive signals in tree-ring carbon and oxygen stable-isotope composition provides increased understanding of how climate and hydrology influenced bog pines (Pinus sylvestris L.) at two sites in southern Sweden during the mid- and late Holocene. Tree-ring sequences from two subfossil trees collected at raised bogs having different hydrology and catchment size were analyzed to probe the stable-isotope signals associated with two bog-wide episodes of growth depression, one during the Holocene Thermal Maximum and the other during the Neoglacial Transition. The occurrence of lower whole-wood δ¹³C and cellulose δ¹³C and δ¹⁸O values immediately prior to the onset of growth depression in both trees, suggesting increased atmospheric relative humidity, is consistent with the notion that excessive effective moisture impeded tree growth. Correlation analysis indicates that the growth response lagged about three years behind the decline in δ¹³C and δ¹⁸O values in each tree, possibly reflecting relatively slow rise in the local water table in response to wetter climate.     

Interpreting species‐specific variation in tree‐ring oxygen isotope ratios among three temperate forest trees

Although considerable variation has been documented in tree‐ring cellulose oxygen isotope ratios (δ¹⁸O_cell) among co‐occurring species, the underlying causes are unknown. Here, we used a combination of field measurements and modelling to investigate the mechanisms behind variations in late‐wood δ¹⁸O_cell (δ¹⁸O_lc) among three co‐occurring species (chestnut oak, black oak and pitch pine) in a temperate forest. For two growing seasons, we quantified among‐species variation in δ¹⁸O_lc, as well as several variables that could potentially cause the δ¹⁸O_lc variation. Data analysis based on the δ¹⁸O_cell model rules out leaf water enrichment (Δ¹⁸O_lw) and tree‐ring formation period (Δt), but highlights source water δ¹⁸O (δ¹⁸O_sw) as an important driver for the measured difference in δ¹⁸O_lc between black and chestnut oak. However, the enriched δ¹⁸O_lc in pitch pine relative to the oaks could not be sufficiently explained by consideration of the above three variables only, but rather, we show that differences in the proportion of oxygen exchange during cellulose synthesis (p_ex) is most likely a key mechanism. Our demonstration of the relevance of some species‐specific features (or lack thereof) to δ¹⁸O_cell has important implications for isotope based ecophysiological/paleoclimate studies.     

The “isohydric trap”: A proposed feedback between water shortage,stomatal regulation,and nutrient acquisition drives differential growth and survival of European pines under climatic dryness

Climatic dryness imposes limitations on vascular plant growth by reducing stomatal conductance, thereby decreasing CO₂ uptake and transpiration. Given that transpiration‐driven water flow is required for nutrient uptake, climatic stress‐induced nutrient deficit could be a key mechanism for decreased plant performance under prolonged drought. We propose the existence of an “isohydric trap,” a dryness‐induced detrimental feedback leading to nutrient deficit and stoichiometry imbalance in strict isohydric species. We tested this framework in a common garden experiment with 840 individuals of four ecologically contrasting European pines (Pinus halepensis, P. nigra, P. sylvestris, and P. uncinata) at a site with high temperature and low soil water availability. We measured growth, survival, photochemical efficiency, stem water potentials, leaf isotopic composition (δ¹³C, δ¹⁸O), and nutrient concentrations (C, N, P, K, Zn, Cu). After 2 years, the Mediterranean species Pinus halepensis showed lower δ¹⁸O and higher δ¹³C values than the other species, indicating higher time‐integrated transpiration and water‐use efficiency (WUE), along with lower predawn and midday water potentials, higher photochemical efficiency, higher leaf P, and K concentrations, more balanced N:P and N:K ratios, and much greater dry‐biomass (up to 63‐fold) and survival (100%). Conversely, the more mesic mountain pine species showed higher leaf δ¹⁸O and lower δ¹³C, indicating lower transpiration and WUE, higher water potentials, severe P and K deficiencies and N:P and N:K imbalances, and poorer photochemical efficiency, growth, and survival. These results support our hypothesis that vascular plant species with tight stomatal regulation of transpiration can become trapped in a feedback cycle of nutrient deficit and imbalance that exacerbates the detrimental impacts of climatic dryness on performance. This overlooked feedback mechanism may hamper the ability of isohydric species to respond to ongoing global change, by aggravating the interactive impacts of stoichiometric imbalance and water stress caused by anthropogenic N deposition and hotter droughts, respectively.