Needle δ13C Research Articles

Seasonal variation in C:N:P stoichiometry, nonstructural carbohydrates, and carbon isotopes of two coniferous pioneer tree species in subtropical China.

The characteristics of C:N:P stoichiometry, nonstructural carbohydrate (NSC) content, and C stable isotopes and their relationships affect plant responses to environmental changes and are critical to understanding the ecosystem carbon and water cycles. We investigated the water use strategies and physiological changes of two pioneer tree species (Pinus armandii and Pinus yunnanensis) in response to seasonal drought in subtropical China. The seasonal variation in needle δ13C values, C:N:P stoichiometry, and NSC contents of the two tree species were studied in 25-year-old plantation in central Yunnan Province. The needle δ13C values of both species were highest in summer. Soluble sugars, starch and NSC content of the two tree species decreased from spring to winter, while there was no significant difference in the seasonal variation of soluble sugars/starch in P. armandii needles, the maximum soluble sugars/starch in P. yunnanensis needles was in autumn. In addition, the C, N, and P contents of the needles and the C:N and C:P ratios of the two species showed different seasonal fluctuations, whereas the N:P ratio decreased with the season. The C:N:P stoichiometry and NSC content of the needles showed significant correlations, whereas the needle δ13C was weakly correlated with C:N:P stoichiometry and NSC content. Phenotypic plasticity analysis and principal component analysis revealed that the needle nutrient characteristics (NSC and P contents and N:P ratio) and needle δ13C values were critical indicators of physiological adaptation strategies of P. armandii and P. yunnanensis for coping with seasonal variation. These results increase our understanding of the water-use characteristics of the two pioneer tree species and the dynamic balance between the NSC, C, N, and P contents of the needles.

Variation of the water-use efficiency and nutrients of Pinus armandii under different ages and growth season

Seasonal changes in carbon (C), nitrogen (N), and phosphorus (P) stoichiometry and water-use efficiency (WUE) in the needles of Pinus armandii were investigated. The needles of P. armandii were sampled in January, May, August, and November 2021. The results showed that (1) the needle δ13C values of young P. armandii forests were higher than those of middle-aged P. armandii forests, and these differences were significant in winter. (2) Both young and middle-aged P. armandii forests were restricted by N and P, and P. armandii was more restricted by N in summer, autumn, and winter. (3) Needle δ13C values were significantly positively correlated with P and negatively correlated with N, C/P, and N/P in young P. armandii forests; P. armandii (especially young stands) can compensate for decreases in N-use efficiency by increasing WUE. The C, N, and P content and their stoichiometric ratios were affected by a combination of season, stand age, and WUE. This study contributes to an in-depth understanding of the adaptive mechanisms of nutrients and WUE in P. armandii in the subtropical region. Our findings also have implications for the management of P. armandii plantations in the study area.

Seasonal variation in δ13C of Pinus. yunnanensis and Pinus. armandii at different stand ages

Seasonal drought is common in Yunnan province, and water is the dominant factor limiting the growth of Pinus. yunnanensis and Pinus. armandii. The water use efficiency (WUE) of the two species is poorly understood. Needles were collected in a plantation (P. yunnanensis and P. armandii mixed forest) in four seasons, and the needle δ13C values were measured. The selected species had larger δ13C values and exhibited higher WUE than typical subtropical species. P. armandii needles showed a more conservative water use strategy (high WUE) than P. yunnanensis. There were significant differences in the δ13C values of P. armandii between the two ages, whereas no difference was observed in the δ13C values of P. yunnanensis. The lowest δ13C value in the young P. armandii forest was observed in spring, whereas the δ13C value of middle-aged forests did not differ between the seasons. The δ13C value of young P. yunnanensis forests showed no difference in the four seasons, and the maximum value was observed in summer in middle-aged forests. In general, the δ13C value of P. armandii was lowest in spring, whereas that of P. yunnanensis was higher in spring and winter. The needle δ13C values were lower in spring and winter, indicating that the season had different effects on the δ13C values of different tree species. Correlation analysis between the needle δ13C values and meteorological data indicated that temperature and precipitation were the dominant factors affecting WUE in P. yunnanensis and P. armandii. The effect of temperature on WUE was greater in P. yunnanensis middle-aged forests. The identification and selection of subtropical tree species with high WUE are critical to maintaining high levels of forest benefits under limited water conditions.

Effects of long-term nitrogen addition on the δ15N and δ13C of Larix gmelinii and soil in a boreal forest

BackgroundNatural abundance of carbon (C) and nitrogen (N) stable isotope ratios (δ13C and δ15N) has been used to indicate the state and cycle of ecosystem C and N. However, it is still unclear how C and N cycle of boreal forests respond to the N deposition.ResultsWe conducted an 8-year continuous N addition field experiment in a Larix gmelinii forest in Greater Khingan Mountains, Northeast China. Four N treatments (0, 25, 50, 75 kg N ha−1 year−1) were built. The effects of N addition on the δ13C and δ15N of needle, branch, bark, and fine root of Larix gmelinii and soil were studied. The result of the balance between the N input and output flux showed that N addition significantly increased the δ15N in each organ of Larix gmelinii, but did not change the δ15N of soil. We also found that the N absorption by needles of Larix gmelinii could increase the needle photosynthesis rate and δ13C by increasing carboxylation, but N addition had no significant effect on the δ13C of soil and other organs. In addition, both the soil δ15N and δ13C increased with the soil depth.ConclusionsLong-term N addition may lead to more open C and N cycles and further affect plant nutrient acquisition strategies in boreal forest ecosystems.

Effect of Drought and Topographic Position on Depth of Soil Water Extraction of Pinus sylvestris L. var. mongolica Litv. Trees in a Semiarid Sandy Region, Northeast China

Drought and topographic position are the most important factors influencing tree growth and survival in semiarid sandy regions of Northeast China. However, little is known about how trees respond to drought in combination with topographic position by modifying the depth of soil water extraction. Therefore, we identified water sources for 33-year-old Mongolian pine (Pinus sylvestris L. var. mongolica Litv.) trees growing at the top and bottom of sand dunes by comparing stable isotopes δ2H and δ18O in twig xylem water, soil water at various depths and groundwater during dry and wet periods. Needle carbon isotope composition (δ13C) was simultaneously measured to assess water use efficiency. Results showed that when soil moisture was low during the dry period, trees at the top used 40–300 cm soil water while trees at the bottom utilized both 40–300 cm soil water and possibly groundwater. Nevertheless, when soil moisture at 0–100 cm depth was higher during the wet period, it was the dominant water sources for trees at both the top and bottom. Moreover, needle δ13C in the dry period were significantly higher than those in the wet period. These findings suggested that trees at both the top and bottom adjust water uptake towards deeper water sources and improve their water use efficiency under drought condition. Additionally, during the dry period, trees at the top used shallower water sources compared with trees at the bottom, in combination with significantly higher needle δ13C, indicating that trees at the bottom applied a relatively more prodigal use of water by taking up deeper water (possibly groundwater) during drought conditions. Therefore, Mongolian pine trees at the top were more susceptible to suffer dieback under extreme dry years because of shallower soil water uptake and increased water restrictions. Nevertheless, a sharp decline in the groundwater level under extreme dry years had a strong negative impact on the growth and survival of Mongolian pine trees at the bottom due to their utilization of deeper water sources (possibly groundwater).

Environmental Controls on the Seasonal Variation in Gas Exchange and Water Balance in a Near-Coastal Mediterranean Pinus halepensis Forest

Aleppo pine (Pinus halepensis Mill.) is widespread in most countries of the Mediterranean area. In Greece, Aleppo pine forms natural stands of high economic and ecological importance. Understanding the species’ ecophysiological traits is important in our efforts to predict its responses to ongoing climate variability and change. Therefore, the aim of this study was to assess the seasonal dynamic in Aleppo pine gas exchange and water balance on the leaf and canopy levels in response to the intra-annual variability in the abiotic environment. Specifically, we assessed needle gas exchange, water potential and δ13C ratio, as well as tree sap flow and canopy conductance in adult trees of a mature near-coastal semi-arid Aleppo pine ecosystem, over two consecutive years differing in climatic conditions, the latter being less xerothermic. Maximum photosynthesis (Amax), stomatal conductance (gs), sap flow per unit leaf area (Ql), and canopy conductance (Gs) peaked in early spring, before the start of the summer season. During summer drought, the investigated parameters were negatively affected by the increasing potential evapotranspiration (PET) rate and vapor pressure deficit (VPD). Aleppo pine displayed a water-saving, drought avoidance (isohydric) strategy via stomatal control in response to drought. The species benefited from periods of high available soil water, during the autumn and winter months, when other environmental factors were not limiting. Then, on the leaf level, air temperature had a significant effect on Amax, while on the canopy level, VPD and net radiation affected Ql. Our study demonstrates the plasticity of adult Aleppo pine in this forest ecosystem in response to the concurrent environmental conditions. These findings are important in our efforts to predict and forecast responses of the species to projected climate variability and change in the region.

Water use strategies of natural Pinus sylvestris var. Mongolica trees of different ages in Hulunbuir Sandy Land of Inner Mongolia, China, based on stable isotope analysis

Natural Mongolian pine trees of different ages consistently use shallow water throughout the main growing season; therefore, water stored in the shallow soil layer is vital for maintaining their viability. Mongolian pine (Pinus sylvestris var. mongolica) plantations in sandy regions often experience dieback after 30–35 years of growth due to water deficiency, whereas natural Mongolian pine forests remain healthy during the same growth stage. However, the water use strategies of natural Mongolian pines remains unclear. Therefore, δ2H and δ18O in twig xylem water, soil water and groundwater were analyzed in 10–20, 20–30 and 30–50-year-old natural Mongolian pine trees to identify their water sources. In addition, needle δ13C was measured simultaneously to assess water use efficiency. Results showed that pine trees of different ages utilized soil water from the same depth. During the growing season (June–August), all pine trees utilized water from 0 to 20 cm soil depth, regardless of the soil water condition. During the end of growing season (September and October), even though soil moisture content in the 0–20 cm depth was higher, pine trees of different ages utilized water from the 0–60 cm soil depth in September and switched to utilize water from the 20–80 cm soil depth in October. There were no significant differences in needle δ13C among the sampling dates for trees in each age group, indicating that pine trees can absorb sufficient water to satisfy their water requirements regardless of age. These findings suggest that water stored in the shallow soil layer (0–20 cm) plays an important role in supporting tree transpiration during the growing season (June–August). Therefore, the stability of shallow soil is vital for maintaining the viability of natural Mongolian pine forests.

Temporal effects of prescribed burning on terpene production in Mediterranean pines.

Prescribed burning is used to reduce fuel hazard but underburning can damage standing trees. The effect of burning on needle terpene storage, a proxy for secondary metabolism, in fire-damaged pines is poorly understood despite the protection terpenes confer against biotic and abiotic stressors. We investigated variation in needle terpene storage after burning in three Mediterranean pine species featuring different adaptations to fire regimes. In two pure-stands of Pinus halepensis Mill. and two mixed-stands of Pinus sylvestris L. and Pinus nigra ssp. salzmanni (Dunal) Franco, we compared 24 h and 1 year post-burning concentrations with pre-burning concentrations in 20 trees per species, and evaluated the relative contribution of tree fire severity and physiological condition (δ13C and N concentration) on temporal terpene dynamics (for mono- sesqui- and diterpenes). Twenty-four hours post-burning, monoterpene concentrations were slightly higher in P. halepensis than at pre-burning, while values were similar in P. sylvestris. Differently, in the more fire-resistant P. nigra monoterpene concentrations were lower at 24 h, compared with pre-burning. One year post-burning, concentrations were always lower compared with pre- or 24 h post-burning, regardless of the terpene group. Mono- and sesquiterpene variations were negatively related to pre-burning δ13C, while diterpene variations were associated with fire-induced changes in needle δ13C and N concentration. At both post-burning times, mono- and diterpene concentrations increased significantly with crown scorch volume in all species. Differences in post-burning terpene contents as a function of the pine species' sensitivity to fire suggest that terpenic metabolites could have adaptive importance in fire-prone ecosystems in terms of flammability or defence against biotic agents post-burning. One year post-burning, our results suggest that in a context of fire-induced resource availability, pines likely prioritize primary rather than secondary metabolism. Overall, this study contributes to the assessment of the direct and indirect effects of fire on pine terpene storage, providing valuable information about their vulnerability to biotic and abiotic stressors throughout time.

Thinning effect on photosynthesis depends on needle ages in a Chinese fir (Cunninghamia lanceolata) plantation.

Canopies in evergreen coniferous plantations often consist of various-aged needles. However, the effect of needle age on the photosynthetic responses to thinning remains ambiguous. Photosynthetic responses of different-aged needles to thinning were investigated in a Chinese fir (Cunninghamia lanceolata) plantation. A dual isotope approach [simultaneous measurements of stable carbon (δ13C) and oxygen (δ18O) isotopes] was employed to distinguish between biochemical and stomatal limitations to photosynthesis. Our results showed that increases in net photosynthesis rates upon thinning only occurred in the current-year and one-year-old needles, and not in the two- to four-year-old needles. The increased δ13C and declined δ18O in current year needles of trees from thinned stands indicated that both the photosynthetic capacity and stomatal conductance resulted in increasing photosynthesis. In one-year-old needles of trees from thinned stands, an increased needle δ13C and a constant needle δ18O were observed, indicating the photosynthetic capacity rather than stomatal conductance contributed to the increasing photosynthesis. The higher water-soluble nitrogen content in current-year and one-year-old needles in thinned trees also supported that the photosynthetic capacity plays an important role in the enhancement of photosynthesis. In contrast, the δ13C, δ18O and water-soluble nitrogen in the two- to four-year-old needles were not significantly different between the control and thinned trees. Thus, the thinning effect on photosynthesis depends on needle age in a Chinese fir plantation. Our results highlight that the different responses of different-aged needles to thinning have to be taken into account for understanding and modelling ecosystem responses to management, especially under the expected environmental changes in future.

Importance of Arboreal Cyanolichen Abundance to Nitrogen Cycling in Sub-Boreal Spruce and Fir Forests of Central British Columbia, Canada

The importance of N2-fixing arboreal cyanolichens to the nitrogen (N)-balance of sub-boreal interior hybrid spruce (Picea glauca × engelmannii) and subalpine fir (Abies lasiocarpa) forests was examined at field sites in central BC, Canada. Host trees were accessed by a single-rope climbing technique and foliage as well as arboreal macrolichen functional groups were sampled by branch height in eight random sample trees from each of two high (High Cyano) and two low (Low Cyano) cyanolichen abundance sites for a total of 32 sample trees. Natural abundances of stable isotopes of N (15N, 14N) and carbon (13C, 12C) were determined for aggregate host tree and epiphytic lichen samples, as well as representative samples of upper organic and soil horizons (Ae and Bf) from beneath host trees. As expected, N2-fixing cyanolichens had 2–6-fold greater N-contents than chlorolichens and a δ15N close to atmospheric N2, while foliage and chlorolichens were more depleted in 15N. By contrast, soils at all trees and sites were 15N-enriched (positive δ15N), with declining (not significant) δ15N with increased tree-level cyanolichen abundance. Lichen functional groups and tree foliage fell into three distinct groups with respect to δ13C; the tripartite cyanolichen Lobaria pulmonaria (lightest), host-tree needles (intermediate), and bipartite cyanolichens, hair (Alectoria and Bryoria spp.) and chlorolichens (heaviest). Branch height of host trees was an effective predictor of needle δ13C. Our results showed a modest positive correlation between host tree foliage N and cyanolichen abundance, supporting our initial hypothesis that higher cyanolichen abundances would elevate host tree foliar N. Further study is required to determine if high cyanolichen abundance enhances host tree and/or stand-level productivity in sub-boreal forests of central BC, Canada.

Comparison of intrinsic water use efficiency between different aged Pinus sylvestris var. mongolica wide windbreaks in semiarid sandy land of northern China

Dieback of Mongolian pine (Pinus sylvestris var. mongolica) wide windbreaks often emerges 35 years after planting in agroforestry systems. Water stress is considered to be the main cause for such dieback, but there is little knowledge about how water use efficiency of trees changes with their age. We measured needle δ13C as a proxy for intrinsic water use efficiency (iWUE), combined with specific leaf area, leaf dry matter content, soil water content, and groundwater level in 9-, 21-, and 41-year-old pine stands in 2011. Results showed that the mean δ13C value of 9-year-old trees was lower by 0.66 and 0.48 ‰ than that of 21- and 41-year-old trees, respectively, but the mean δ13C value in 21-year-old trees was higher by 0.17 ‰ than that in 41-year-old trees. Soil water content for 9-year-old trees was highest. Specific leaf area for 21-year-old trees was lowest. Compared with the other two tree ages, tree roots reached groundwater levels for 41-year-old trees. These results suggested that (1) 9-year-old trees with the lowest iWUE applied a prodigal water use strategy due to higher soil moisture, (2) 21-year-old trees with the highest iWUE employed a more conservative water use strategy and experienced significant water stress because of lower soil moisture and specific leaf area, and (3) the water use strategy of 41-year-old trees was between that of 9- and 21-year-old trees, likely due to both aging and uptake of groundwater. Dieback of Mongolian pine trees at older age (e.g., more than 35 years old) might occur when the groundwater level declines suddenly in years of extremely low rainfall.

Importance of soil moisture and N availability to larch growth and distribution in the Arctic taiga-tundra boundary ecosystem, northeastern Siberia

To better understand the factors controlling the growth of larch trees in Arctic taiga-tundra boundary ecosystem, we conducted field measurements of photosynthesis, tree size, nitrogen (N) content, and isotopic ratios in larch needles and soil. In addition, we observed various environmental parameters, including topography and soil moisture at four sites in the Indigirka River Basin, near Chokurdakh, northeastern Siberia. Most living larch trees grow on mounds with relatively high elevations and dry soils, indicating intolerance of high soil moisture. We found that needle δ13C was positively correlated with needle N content and needle mass, and these parameters showed spatial patterns similar to that of tree size. These results indicate that trees with high needle N content achieved higher rates of photosynthesis, which resulted in larger amounts of C assimilation and larger C allocation to needles and led to larger tree size than trees with lower needle N content. A positive correlation was also found between needle N content and soil NH4+ pool. Thus, soil inorganic N pool may indicate N availability, which is reflected in the needle N content of the larch trees. Microtopography plays a principal role in N availability, through a change in soil moisture. Relatively dryer soil of mounds with higher elevation and larger extent causes higher rates of soil N production, leading to increased N availability for plants, in addition to larger rooting space for trees to uptake more N.

Needle δ13C and mobile carbohydrates in Pinus koraiensis in relation to decreased temperature and increased moisture along an elevational gradient in NE China

A tree’s crown interacts with atmospheric variables such as CO2, temperature, and humidity. Physioecology of leaves/needles (e.g. δ13C, mobile carbohydrates, and nitrogen) is, therefore, strongly affected by microclimate in and surrounding a tree crown. To understand the physiological responses of leaves to changes in air temperature and moisture, we measured δ13C, soluble sugars, starch, and total nitrogen (N) concentrations in current year and 1-yr-old needles of Pinus koraiensis trees, and compared the growing season air temperature and relative humidity within and outside P. koraiensis crowns along an elevational gradient from 760 to 1,420 m a.s.l. on Changbai Mountain, NE China. Our results indicated that needle N and mobile carbohydrates concentrations, as well as needle δ13C values changed continuously with increasing elevation, corresponding to a continuous decrease in air temperature and an increase in relative humidity. Needle carbon and nitrogen status is highly significantly negatively correlated with temperature, but positively correlated with relative humidity. These results indicate that increases in air temperature in combination with decreases in relative humidity may result in lower levels of N and mobile carbohydrates in P. koraiensis trees, suggesting that future climate changes such as global warming and changes in precipitation patterns will directly influence the N and carbon physiology at P. koraiensis individual level, and indirectly affect the competitive ability, species composition, productivity and functioning at the stand and ecosystem level in NE China. Due to the relatively limited range of the transect (760–1,420 m) studied, further research is needed to explain whether the present results are applicable to scales across large elevational gradients.

Needle-age related variability in nitrogen, mobile carbohydrates, and δ13C within Pinus koraiensis tree crowns.

For both ecologists and physiologists, foliar physioecology as a function of spatially and temporally variable environmental factors such as sunlight exposure within a tree crown is important for understanding whole tree physiology and for predicting ecosystem carbon balance and productivity. Hence, we studied concentrations of nitrogen (N), non-structural carbohydrates (NSC = soluble sugars + starch), and δ13C in different-aged needles within Pinus koraiensis tree crowns, to understand the needle age- and crown position-related physiology, in order to test the hypothesis that concentrations of N, NSC, and δ13C are needle-age and crown position dependent (more light, more photosynthesis affecting N, NSC, and δ13C), and to develop an accurate sampling strategy. The present study indicated that the 1-yr-old needles had significantly higher concentration levels of mobile carbohydrates (both on a mass and an area basis) and Narea (on an area basis), as well as NSC-N ratios, but significantly lower levels of Nmass (on a mass basis) concentration and specific leaf area (SLA), compared to the current-year needles. Azimuthal (south-facing vs. north-facing crown side) effects were found to be significant on starch [both on a mass (STmass) and an area basis (STarea)], δ13C values, and Narea, with higher levels in needles on the S-facing crown side than the N-facing crown side. Needle Nmass concentrations significantly decreased but needle STmass, STarea, and δ13C values significantly increased with increasing vertical crown levels. Our results suggest that the sun-exposed crown position related to photosynthetic activity and water availability affects starch accumulation and carbon isotope discrimination. Needle age associated with physiological activity plays an important role in determining carbon and nitrogen physiology. The present study indicates that across-scale sampling needs to carefully select tissue samples with equal age from a comparable crown position.

The effects of Arceuthobium sichuanense infection on needles and current‐year shoots of mature and young Qinghai spruce (Picea crassifolia) trees

SummaryArceuthobium sichuanense is a hemiparasitic angiosperm that infects Qinghai spruce (Picea crassifolia Kom.) in Qinghai province, China, and causes severe damage to spruce forests in Qinghai‐Tibet Plateau. In this study, the impact of A. sichuanense infection on mature and young trees of Qinghai spruce was evaluated by examining needle and current‐year shoot morphology, needle water and nitrogen‐use efficiency (NUE) and needle nitrogen concentration. The most apparent effect of A. sichuanense infection was a significant reduction in both needle size distal to infection and current‐year shoot length in the infected branches (p < 0.001). Per cent reductions in needle and current‐year shoot length were similar between mature and young trees (58.9 vs. 56.3%; 59.7 vs. 62.9%). There was a high degree of correlation in foliar δ15N values between the dwarf mistletoe and its host trees (R2 = 0.9017, p < 0.001), while the foliar δ13C values of A. sichuanense were similar to those of infected mature and young spruce trees. The dwarf mistletoe infection also resulted in a significant decrease in host needle N concentration and δ13C values (p < 0.001). The per cent reduction in needle N concentration in young trees was nearly twice as much as that in mature trees (20.49 vs. 11.54%), while the per cent reduction in needle δ13C values was similar between young and mature trees (−0.98 vs.−1.1‰). The NUE in mature trees was not affected by A. sichuanense infection, but the NUE in young trees was increased by the infection.

From pattern to process: linking intrinsic water‐use efficiency to drought‐induced forest decline

AbstractThe rise in atmospheric CO2 concentrations (Ca) has been related to tree growth enhancement and increasing intrinsic water‐use efficiency (iWUE). However, the extent that rising Ca has led to increased long‐term iWUE and whether climate could explain deviations from expected Ca‐induced growth enhancement are still poorly understood. The aim of this research was to use Ca and local climatic variability to explain changes during the 20th century in growth and tree ring and needle δ13C in declining and nondeclining Abies alba stands from the Spanish Pyrenees, near the southern distribution limit of this species. The temporal trends of iWUE were calculated under three theoretical scenarios for the regulation of plant‐gas exchange at increasing Ca. We tested different linear mixed‐effects models by multimodel selection criteria to predict basal area increment (BAI), a proxy of tree radial growth, using these scenarios and local temperature together with precipitation data as predictors. The theoretical scenario assuming the strongest response to Ca explained 66–81% of the iWUE variance and 28–56% of the observed BAI variance, whereas local climatic variables together explained less than 11–21% of the BAI variance. Our results are consistent with a drought‐induced limitation of the tree growth response to rising CO2 and a decreasing rate of iWUE improvement from the 1980s onward in declining A. alba stands subjected to lower water availability.

Association between tree-ring and needle δ13C and leaf gas exchange in Pinus halepensis under semi-arid conditions

Association between tree-ring and needle δ13C and leaf gas exchange in Pinus halepensis under semi-arid conditions

Association between tree-ring and needle δ13C and leaf gas exchange in Pinus halepensis under semi-arid conditions

Association between tree-ring and needle δ13C and leaf gas exchange in Pinus halepensis under semi-arid conditions

Association between tree-ring and needle δ13C and leaf gas exchange in Pinus halepensis under semi-arid conditions

Association between tree-ring and needle δ13C and leaf gas exchange in Pinus halepensis under semi-arid conditions

Association between tree-ring and needle δ13C and leaf gas exchange in Pinus halepensis under semi-arid conditions

Association between tree-ring and needle δ13C and leaf gas exchange in Pinus halepensis under semi-arid conditions