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1.
Canopy CO2 concentrations in a tropical rainforest in French Guiana were measured continuously for 5 days during the 1994 dry season and the 1995 wet season. Carbon dioxide concentrations ([CO2]) throughout the canopy (0.02–38 m) showed a distinct daily pattern, were well-stratified and decreased with increasing height into the canopy. During both seasons, daytime [CO2] in the upper and middle canopy decreased on average 7–10 μmol mol−1 below tropospheric baseline values measured at Barbados. Within the main part of the canopy (≥ 0.7 m), [CO2] did not differ between the wet and dry seasons. In contrast, [CO2] below 0.7 m were generally higher during the dry season, resulting in larger [CO2] gradients. Supporting this observation, soil CO2 efflux was on average higher during the dry season than during the wet season, either due to diffusive limitations and/or to oxygen deficiency of root and microbial respiration. Soil respiration rates decreased by 40% after strong rain events, resulting in a rapid decrease in canopy [CO2] immediately above the forest floor of about 50␣μmol mol−1. Temporal and spatial variations in [CO2]canopy were reflected in changes of δ13Ccanopy and δ18Ocanopy values. Tight relationships were observed between δ13C and δ18O of canopy CO2 during both seasons (r 2 > 0.86). The most depleted δ13Ccanopy and δ18Ocanopy values were measured immediately above the forest floor (δ13C = −16.4‰; δ18O = 39.1‰ SMOW). Gradients in the isotope ratios of CO2 between the top of the canopy and the forest floor ranged between 2.0‰ and 6.3‰ for δ13C, and between 1.0‰ and 3.5‰ for δ18O. The δ13Cleaf and calculated c i/c a of foliage at three different positions were similar for the dry and wet seasons indicating that the canopy maintained a constant ratio of photosynthesis to stomatal conductance. About 20% of the differences in δ13Cleaf within the canopy was accounted for by source air effects, the remaining 80% must be due to changes in c i/c a. Plotting 1/[CO2] vs. the corresponding δ13C ratios resulted in very tight, linear relationships (r 2 = 0.99), with no significant differences between the two seasons, suggesting negligible seasonal variability in turbulent mixing relative to ecosystem gas exchange. The intercepts of these relationships that should be indicative of the δ13C of respired sources were close to the measured δ13C of soil respired CO2 and to the δ13C of litter and soil organic matter. Estimates of carbon isotope discrimination of the entire ecosystem, Δe, were calculated as 20.3‰ during the dry season and as 20.5‰ during the wet season. Received: 3 March 1996 / Accepted: 19 October 1996  相似文献   

2.
In this study, sun leaf carbon isotope composition (δ13C) of two co-occurring woody Mediterranean species (Quercus pubescens Willd., a deciduous oak, and Q. ilex L., an evergreen one) was investigated on four sites with different water availability. The total range of δ13C values was 4.4 and 3.1‰ for Q. pubescens and Q. ilex respectively. The intra-site variability was about 3‰. Total mean per species was equal. There were significant differences among sites, but at each site means of δ13C were not significantly different between species. A simple physiological model predicts no difference in intrinsic water-use efficiency (WUEi) between evergreen and deciduous oaks. The relationship between site means of δ13C and water parameters suggests that there is a leaf functional adjustment with respect to available water resource. No correlation was found between δ13C and the contents of any mass-based biochemical constituent. Nevertheless there was a significant correlation between δ13C and leaf mass per area of Q. ilex. For both species, there is also a positive correlation between leaf δ13C and individual crown area, i.e. a structural characteristic at tree level. Causal relations between δ13C and plant-environment interactions are discussed. Received: 25 October 1996 / Accepted: 19 January 1997  相似文献   

3.
The vertical profile of stable carbon isotope ratios (δ13C) of leaves was analyzed for 13 tree species in a cool-temperate deciduous forest in Japan. The vertical distribution of long-term averaged δ13C in atmospheric CO2a) was estimated from δ13C of dry matter from NADP-malic enzyme type C4 plant (Zea mays L. var. saccharata Sturt.) grown at a tower in the forest for 32␣days, assuming constant Δ value (3.3‰) in Z. mays against height. The δa value obtained from δ13C in Z.␣mays was lowest at the forest floor (−9.30 ± 0.03‰), increased with height, and was almost constant above 10␣m (−7.14 ± 0.14‰). Then leaf Δ values for the tree species were calculated from tree leaf δ13 C andδa. Mean leaf Δ values for the three tall deciduous species (Fraxinus mandshurica, Ulmus davidiana, and Alnus hirsuta) were significantly different among three height levels in the forest: 23.1 ± 0.7‰ at the forest floor (understory), 21.4 ± 0.5‰ in lower canopy, and 20.5 ± 0.3‰ in upper canopy. The true difference in tree leaf Δ among the forest height levels might be even greater, because Δ in Z. mays probably increased with shading by up to ∼‰. The difference in tree leaf Δ among the forest height levels would be mainly due to decreasing intercellular CO2 (C i) with the increase in irradiance. Potential assimilation rate for the three tree species probably increased with height, since leaf nitrogen content on an area basis for these species also increased with height. However, the increase in stomatal conductance for these tree species would fail to meet the increase in potential assimilation rate, which might lead to increasing the degree of stomatal limitation in photosynthesis with height. Received: 30 September 1995 / Accepted: 25 October 1996  相似文献   

4.
We tested the hypothesis that elevated CO2 would stimulate proportionally higher photosynthesis in the lower crown of Populus trees due to less N retranslocation, compared to tree crowns in ambient CO2. Such a response could increase belowground C allocation, particularly in trees with an indeterminate growth pattern such as Populus tremuloides. Rooted cuttings of P. tremuloides were grown in ambient and twice ambient (elevated) CO2 and in low and high soil N availability (89 ± 7 and 333 ± 16 ng N g−1 day−1 net mineralization, respectively) for 95 days using open-top chambers and open-bottom root boxes. Elevated CO2 resulted in significantly higher maximum leaf photosynthesis (A max) at both soil N levels. A max was higher at high N than at low N soil in elevated, but not ambient CO2. Photosynthetic N use efficiency was higher at elevated than ambient CO2 in both soil types. Elevated CO2 resulted in proportionally higher whole leaf A in the lower three-quarters to one-half of the crown for both soil types. At elevated CO2 and high N availability, lower crown leaves had significantly lower ratios of carboxylation capacity to electron transport capacity (V cmax/J max) than at ambient CO2 and/or low N availability. From the top to the bottom of the tree crowns, V cmax/J max increased in ambient CO2, but it decreased in elevated CO2 indicating a greater relative investment of N into light harvesting for the lower crown. Only the mid-crown leaves at both N levels exhibited photosynthetic down regulation to elevated CO2. Stem biomass segments (consisting of three nodes and internodes) were compared to the total A leaf for each segment. This analysis indicated that increased A leaf at elevated CO2 did not result in a proportional increase in local stem segment mass, suggesting that C allocation to sinks other than the local stem segment increased disproportionally. Since C allocated to roots in young Populus trees is primarily assimilated by leaves in the lower crown, the results of this study suggest a mechanism by which C allocation to roots in young trees may increase in elevated CO2. Received: 12 August 1996 / Accepted: 12 November 1996  相似文献   

5.
Carbon isotope composition of boreal plants: functional grouping of life forms   总被引:22,自引:0,他引:22  
 We tested the hypothesis that life forms (trees, shrubs, forbs, and mosses; deciduous or evergreen) can be used to group plants with similar physiological characteristics. Carbon isotope ratios (δ13C) and carbon isotope discrimination (Δ) were used as functional characteristics because δ13C and Δ integrate information about CO2 and water fluxes, and so are useful in global change and scaling studies. We examined δ13C values of the dominant species in three boreal forest ecosystems: wet Picea mariana stands, mesic Populus tremuloides stands, and dry Pinus banksiana stands. Life form groups explained a significant fraction of the variation in leaf carbon isotope composition; seven life-form categories explained 50% of the variation in δ13C and 42% of the variation in Δ and 52% of the variance not due to intraspecific genetic differences (n=335). The life forms were ranked in the following order based on their values: evergreen trees<deciduous trees=evergreen and deciduous shrubs=evergreen forbs<deciduous forbs=mosses. This ranking of the life forms differed between deciduous (Populus) and evergreen (Pinus and Picea) ecosystems. Furthermore, life forms in the Populus ecosystem had higher discrimination values than life forms in the dry Pinus ecosystem; the Picea ecosystem had intermediate Δ values. These correlations between Δ and life form were related to differences in plant stature and leaf longevity. Shorter plants had lower Δ values than taller plants, resulting from reduced light intensity at lower levels in the forest. After height differences were accounted for, deciduous leaves had higher discrimination values than evergreen leaves, indicating that deciduous leaves maintained higher ratios of intracellular to ambient CO2 (c i/c a) than did evergreen leaves in a similar environment within these boreal ecosystems. We found the same pattern of carbon isotope discrimination in a year with above-average precipitation as in a year with below-average precipitation, indicating that environmental fluctuations did not affect the ranking of life forms. Furthermore, plants from sites near the northern and southern boundaries of the boreal forest had similar patterns of discrimination. We concluded that life forms are robust indicators of functional groups that are related to carbon and water fluxes within boreal ecosystems. Received: 15 April 1996 / Accepted: 16 November 1996  相似文献   

6.
The interspecific variability of sunlit leaf carbon isotope composition (δ13C), an indicator of leaf intrinsic water-use efficiency (WUE, CO2 assimilation rate/leaf conductance for water vapour), was investigated in canopy trees of three lowland rainforest stands in French Guiana, differing in floristic composition and in soil drainage characteristics, but subjected to similar climatic conditions. We sampled leaves with a rifle from 406 trees in total, representing 102 species. Eighteen species were common to the three stands. Mean species δ13C varied over a 6.0‰ range within each stand, corresponding to WUE varying over about a threefold range. Species occurring in at least two stands displayed remarkably stable δ13C values, suggesting a close genetic control of species δ13C. Marked differences in species δ13C values were found with respect to: (1) the leaf phenology pattern (average δ13C=–29.7‰ and –31.0‰ in deciduous-leaved and evergreen-leaved species, respectively), and (2) different types of shade tolerance defined by features reflecting the plasticity of growth dynamics with respect to contrasting light conditions. Heliophilic species exhibited more negative δ13C values (average δ13C=–30.5‰) (i.e. lower WUE) than hemitolerant species (–29.3‰). However, tolerant species (–31.4‰) displayed even more negative δ13C values than heliophilic ones. We could not provide a straightforward ecophysiological interpretation of this result. The negative relationship found between species δ13C and midday leaf water potential (Ψwm) suggests that low δ13C is associated with high whole tree leaf specific hydraulic conductance. Canopy carbon isotope discrimination (Δ A ) calculated from the basal area-weighed integral of the species δ13C values was similar in the three stands (average Δ A =23.1‰), despite differences in stand species composition and soil drainage type, reflecting the similar proportions of the three different shade-tolerance types among stands. Received: 30 November 1999 / Accepted: 23 March 2000  相似文献   

7.
The natural ratio of stable carbon isotopes (δ13C) was compared to leaf structural and chemical characteristics in evergreen conifers in the north-central Rockies, United States. We sought a general model that would explain variation in δ13C across altitudinal gradients. Because variation in δ13C is attributed to the shifts between supply and demand for carbon dioxide within the leaf, we measured structural and chemical variables related to supply and demand. We measured stomatal density, which is related to CO2 supply to the chloroplasts, and leaf nitrogen content, which is related to CO2 demand. Leaf mass per area was measured as an intermediate between supply and demand. Models were tested on four evergreen conifers: Pseudotsuga menziesii, Abies lasiocarpa, Picea engelmannii, and Pinus contorta, which were sampled across 1800 m of altitude. We found significant variation among species in the rate of δ13C increase with altitude, ranging from 0.91‰ km–1 for A. lasiocarpa to 2.68‰ km–1 for Pinus contorta. Leaf structure and chemistry also varied with altitude: stomatal density decreased, leaf mass per area increased, but leaf nitrogen content (per unit area) was constant. The regressions on altitude were particularly robust in Pinus contorta. Variables were derived to describe the balance between supply and demand; these variables were stomata per gram of nitrogen and stomata per gram of leaf mass. Both derived variables should be positively related to internal CO2 supply and thus negatively related to δ13C. As expected, both derived variables were negatively correlated with δ13C. In fact, the regression on stomatal density per gram was the best fit in the study (r 2=0.72, P<0.0001); however, the relationships were species specific. The only general relationship observed was between δ13C and LMA: δ13C (‰)=–32.972+ 0.0173×LMA (r 2=0.45, P<0.0001). We conclude that species specificity of the isotopic shift indicates that evergreen conifers demonstrate varying degrees of functional plasticity across environmental gradients, while the observed convergence of δ13C with LMA suggests that internal resistance may be the key to understanding inter-specific isotopic variation across altitude. Received: 1 June 1999 / Accepted: 2 November 1999  相似文献   

8.
Vertical CO2 profiles (between 0.02 and 14.0 m) were studied in forest canopies of Pinus contorta, Populus tremuloides, and in a riparian forest with Acer negundo and Acer grandidentatum during two consecutive growing seasons. Profiles, measured continuously during 1- to 13-day periods in four to five stands differing in overstorey canopy area index (CAI < 4.5; including leaves, branches and stems), were well stratified, with highest [CO2] just above the forest floor. Canopy [CO2] profiles were influenced by stand structure (CAI, presence of understorey vegetation), and were highly dependent on vegetation type (deciduous and evergreen). A doubling of CAI in Acer spp. and P. tremuloides stands did not show an effect on upper canopy [CO2], when turbulent mixing was high. However, increasing understorey biomass in Acer spp. stands had a profound effect on lower canopy [CO2]. In open stands with a vigorous understorey layer, higher soil respiration rates were offset by increased understorey gas exchange, resulting in [CO2] below those of the convective boundary layer (CBL). Midday depletions up to 20 ppmv below CBL values could be frequently observed in deciduous canopies. In evergreen canopies, [CO2] stayed generally above the CBL background values, [CO2] profiles were more uniform, and gradients were smaller than in deciduous stands with similar CAI. Seasonal changes of canopy [CO2] reflected changes in soil respiration rates as well as plant phenology and gas exchange of both dominant tree and understorey vegetation. Seasonal patterns were less pronounced in evergreen than in deciduous forests.  相似文献   

9.
A. Ares  J. H. Fownes 《Oecologia》1999,121(4):458-466
We studied changes in stand structure, productivity, canopy development, growth efficiency, and intrinsic water use efficiency (WUE=photosynthesis/stomatal conductance) of the native tree koa (Acacia koa) across a gradient of decreasing rainfall (2600–700 mm) with increasing elevation (700–2000 m) on the island of Hawaii. The stands were located on organic soils on either smooth (pahoehoe) or rough (aa) lava flows. In the greenhouse, we also examined growth and WUE responses to different water regimes of koa seedlings grown from seeds collected in the study area. We tested the hypotheses that (1) stand basal area, aboveground net primary productivity (ANPP), leaf area index (LAI), and growth per unit leaf area decreased with decreasing rainfall, (2) WUE increased with decreasing rainfall or water supply, and (3) WUE responses were caused by stomatal limitation rather than by nutrient limitations to photosynthesis. The carbon isotope composition of phyllode tissues (δ13C) was examined as an integrated measure of WUE. Basal area and LAI of koa stands on both pahoehoe and aa lava flows, and ANPP on aa lava flows decreased with elevation. Basal area, LAI, and ANPP of koa in mixed stands with the exotic tropical ash (Fraxinus udhei) were lower compared to single-species koa stands at similar elevations. Along the gradient, phyllode δ13C (and therefore WUE) increased with elevation from –30.2 to –26.8‰. Koa in mixed stands exhibited higher (less negative) δ13C than in single-species stands suggesting that koa and tropical ash competed for water. In the greenhouse, we observed the same trend observed in the field, as phyllode δ13C increased from –27.7 to –24‰ as water supply decreased. Instantaneous gas exchange measurements in the greenhouse showed an inverse correlation of both maximum (morning) photosynthesis (A) and conductance (g) with δ13C values and, also, a good agreement between instantaneous (A/g) and integrated measures of WUE. Phyllode δ13C was not correlated with foliar concentrations of N or other nutrients in either the field or the greenhouse, indicating that differences in δ13C were caused by stomatal limitation rather than by nutrient-related changes in photosynthetic capacity. This study provided evidence that long-term structural and growth adjustments as well as changes in WUE are important mechanisms of koa response to water limitation.  相似文献   

10.
This study investigated the effects on running economy (RE) of ingesting either no fluid or an electrolyte solution with or without 6% carbohydrate (counterbalanced design) during 60-min running bouts at 80% maximal oxygen consumption (O2max). Tests were undertaken in either a thermoneutral (22–23°C; 56–62% relative humidity, RH) or a hot and humid natural environment (Singapore: 25–35°C; 66–77% RH). The subjects were 15 young adult male Singaporeans [O2max = 55.5 (4.4 SD) ml kg−1 min−1]. The RE was measured at 3 m s−1 [65 (6)% O2max] before (RE1) and after each prolonged run (RE2). Fluids were administered every 2 min, at an individual rate determined from prior tests, to maintain body mass (group mean = 17.4 ml min−1). The O2 during RE2 was higher (P < 0.05) than that during the RE1 test for all treatments, with no differences between treatments (ANOVA). The mean increase in O2 from RE1 to RE2 ranged from 3.4 to 4.7 ml kg−1 min−1 across treatments. In conclusion, the deterioration in RE at 3 m s−1 (65% O2max) after 60 min of running at 80% O2max appears to occur independently of whether fluid is ingested and regardless of whether the fluid contains carbohydrates or electrolytes, in both a thermoneutral and in a hot, humid environment. Accepted: 30 October 1997  相似文献   

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