Foliar Delta Research Articles

Isotopic evidence for oligotrophication of terrestrial ecosystems.

Human societies depend on an Earth system that operates within a constrained range of nutrient availability, yet the recent trajectory of terrestrial nitrogen (N) availability is uncertain. Examining patterns of foliar N concentrations and isotope ratios (δ15N) from more than 43,000 samples acquired over 37 years, here we show that foliar N concentration declined by 9% and foliar δ15N declined by 0.6-1.6‰. Examining patterns across different climate spaces, foliar δ15N declined across the entire range of mean annual temperature and mean annual precipitation tested. These results suggest declines in N supply relative to plant demand at the global scale. In all, there are now multiple lines of evidence of declining N availability in many unfertilized terrestrial ecosystems, including declines in δ15N of tree rings and leaves from herbarium samples over the past 75-150 years. These patterns are consistent with the proposed consequences of elevated atmospheric carbon dioxide and longer growing seasons. These declines will limit future terrestrial carbon uptake and increase nutritional stress for herbivores.

Divergent patterns of foliar 13C and 15N in Quercus aquifolioides with an altitudinal transect on the Tibetan Plateau: an integrated study based on multiple key leaf functional traits

Aims With a close association with plant water availability, foliar delta C-13 had been investigated extensively in alpine regions; however, foliar delta N-15 has rarely been concurrently used as an indicator of plant nitrogen availability. Due to the positive correlations between leaf nitrogen content and foliar delta C-13 and delta N-15 found in previous studies, we expected that they should show consistent patterns along an altitudinal gradient. Methods To test our hypothesis, we measured foliar delta C-13 and delta N-15 in conjunction with multiple key leaf functional traits of Quercus aquifolioides, a dominant species of alpine forest on the eastern slopes of the Sygera Mountains, southeastern Tibetan Plateau from 2500 to 3800 m. Important findings (i) Contrary to our hypothesis, foliar delta C-13 exhibited a significant positive linear relationship with altitude; in contrast, foliar delta N-15 initially increased and subsequently decreased with altitude, the change in trend occurring around 3300 m. (ii) Our analyses indicated that leaf internal resistance and stomatal conductance, rather than photosynthetic capacity indicated by leaf N concentration, apparently explained the altitudinal variation in foliar delta C-13, while differences in foliar delta N-15 were likely the result of soil N availability. (iii) Principal component analysis revealed a clear association between delta C-13 and a tradeoff between water loss and carbon gain, indicated by traits related to gas exchange such as leaf thickness, density, stomatal properties. In contrast, the second axis was associated with delta N-15 and nitrogen acquisition strategy in Q. aquifolioides across its altitudinal distribution, represented by traits related to nitrogen concentration and stomata per gram of leaf nitrogen.

Nitrogen niches revealed through species and functional group removal in a boreal shrub community

Most theories attempting to explain the coexistence of species in local communities make fundamental assumptions regarding whether neighbors exhibit competitive, neutral, or positive resource-use interactions; however, few long-term data from naturally assembled plant communities exist to test these assumptions. We utilized a 13-year experiment consisting of factorial removal of three shrub species (Vaccinium myrtillus, V. vitis-idaea, and Empetrum hermaphroditum) and factorial removal of two functional groups (tree roots and feather mosses) to assess how neighbors affect N acquisition and growth of each of the three shrub species. The removal plots were established on each of 30 lake islands in northern Sweden that form a natural gradient of resource availability. We tested the hypotheses that: (1) the presence of functionally similar neighbors would reduce shrub N acquisition through competition for a shared N resource; (2) the removal of functional groups would affect shrub N acquisition by altering the breadth of their niches; and (3) soil fertility would influence the effects of neighbor removals. We found that the removal of functionally similar neighbors (i.e., other shrub species) usually resulted in higher biomass and biomass N, with the strength of these effects varying strongly with site fertility. Shrub species removals never resulted in altered stable N isotope ratios (delta(15)N), suggesting that the niche breadth of the three shrubs was unaffected by the presence of neighboring shrub species. In the functional group removal experiment, we found positive effects of feather moss removal on V. myrtillus biomass and biomass N, and negative effects on E. hermaphrotium N concentration and V. vitis-idaea biomass and biomass N. Tree root removal also caused a significant shift in foliar delta(15)N of V. myrtillus and altered the delta(15)N, biomass, and biomass N of E. hermaphroditum. Collectively, these results show that the resource acquisition and niche breadth of the three shrub species are often affected by neighbors, and further that both the identity of neighbors and site fertility strongly determine whether these interactions are positive, negative, or neutral. These findings have implications for understanding species coexistence and the reciprocal relationships between productivity and species diversity in this ecosystem.

Patterns and driving factors of WUE and NUE in natural forest ecosystems along the North-South Transect of Eastern China

From July 2008 to August 2008, 72 leaf samples from 22 species and 81 soil samples in the nine natural forest ecosystems were collected, from north to south along the North-South Transect of Eastern China (NSTEC). Based on these samples, we studied the geographical distribution patterns of vegetable water use efficiency (WUE) and nitrogen use efficiency (NUE), and analyzed their relationship with environmental factors. The vegetable WUE and NUE were calculated through the measurement of foliar delta(13)C and C/N of predominant species, respectively. The results showed: (1) vegetable WUE, ranging from 2.13 to 28.67 mg C g(-1) H(2)O, increased linearly from south to north in the representative forest ecosystems along the NSTEC, while vegetable NUE showed an opposite trend, increasing from north to south, ranging from 12.92 to 29.60 g C g(-1) N. (2) Vegetable WUE and NUE were dominantly driven by climate and significantly affected by soil nutrient factors. Based on multiple stepwise regression analysis, mean annual temperature, soil phosphorus concentration, and soil nitrogen concentration were responding for 75.5% of the variations of WUE (p<0.001). While, mean annual precipitation and soil phosphorus concentration could explain 65.7% of the change in vegetable NUE (p<0.001). Moreover, vegetable WUE and NUE would also be seriously influenced by atmospheric nitrogen deposition in nitrogen saturated ecosystems. (3) There was a significant trade-off relationship between vegetable WUE and NUE in the typical forest ecosystems along the NSTEC (p<0.001), indicating a balanced strategy for vegetation in resource utilization in natural forest ecosystems along the NSTEC. This study suggests that global change would impact the resource use efficiency of forest ecosystems. However, vegetation could adapt to those changes by increasing the use efficiency of shortage resource while decreasing the relatively ample one. But extreme impacts, such as heavy nitrogen deposition, would break this trade-off mechanism and give a dramatic disturbance to the ecosystem biogeochemical cycle.

The natural abundance of 15N in plant and soil‐available N indicates a shift of main plant N resources to NO from NH along the N leaching gradient

To investigate which of ammonium (NH(4)(+)) or nitrate (NO(3)(-)) is used by plants at gradient sites with different nitrogen (N) availability, we measured the natural abundance of (15)N in foliage and soil extractable N. Hinoki cypress (Chamaecyparis obtusa Endlicher) planted broadly in Japan was selected for use in this study. We estimated the source proportion of foliar N (NH(4)(+) vs. NO(3)(-)) quantitatively using mass balance equations. The results showed that C. obtusa used mainly NH(4)(+) in N-limited forests, although the dependence of C. obtusa on NO(3)(-) was greater in other NO(3)(-)-rich forests. We regarded dissolved organic N (DON) as a potential N source because a previous study demonstrated that C. obtusa can take up glycine. Thus we added DON to our mass balance equations and calculated the source proportion using an isotope-mixing model (IsoSource model). The results still showed a positive correlation between the calculated plant N proportion of NO(3)(-) and the NO(3)(-) pool size in the soil, indicating that high NO(3)(-) availability increases the reliance of C. obtusa on NO(3)(-). Our data suggest the shift of the N source for C. obtusa from NH(4)(+) to NO(3)(-) according to the relative availability of NO(3)(-). They also show the potential of the foliar delta(15)N of C. obtusa as an indicator of the N status in forest ecosystems with the help of the delta(15)N values of soil inorganic and organic N.

Nitrogen fertilization enhances water-use efficiency in a saline environment

Effects of salinity and nutrients on carbon gain in relation to water use were studied in the grey mangrove, Avicennia marina, growing along a natural salinity gradient in south-eastern Australia. Tall trees characterized areas of seawater salinities (fringe zone) and stunted trees dominated landward hypersaline areas (scrub zone). Trees were fertilized with nitrogen (+N) or phosphorus (+P) or unfertilized. There was no significant effect of +P on shoot growth, whereas +N enhanced canopy development, particularly in scrub trees. Scrub trees maintained greater CO(2) assimilation per unit water transpired (water-use efficiency, WUE) and had lower nitrogen-use efficiency (NUE; CO(2) assimilation rate per unit leaf nitrogen) than fringe trees. The CO(2) assimilation rates of +N trees were similar to those in other treatments, but were achieved at lower transpiration rates, stomatal conductance and intercellular CO(2) concentrations. Maintaining comparable assimilation rates at lower stomatal conductance requires greater ribulose 1.5-bisphosphate carboxylase/oxygenase activity, consistent with greater N content per unit leaf area in +N trees. Hence, +N enhanced WUE at the expense of NUE. Instantaneous WUE estimates were supported by less negative foliar delta(13)C values for +N trees and scrub control trees. Thus, nutrient enrichment may alter the structure and function of mangrove forests along salinity gradients.

Bulk leaf δ18O and δ13C reflect the intensity of intraspecific competition for water in a semi‐arid tussock grassland

We investigated the extent to which plant water and nutrient status are affected by intraspecific competition intensity and microsite quality in a monodominant tussock grassland. Leaf gas exchange and stable isotope measurements were used to assess the water relations of Stipa tenacissima tussocks growing along a gradient of plant cover and soil depth in a semi-arid catchment of Southeast Spain. Stomatal conductance and photosynthetic rate decreased with increasing intensity of competition during the wet growing season, leading to foliar delta(18)O and delta(13)C enrichment. A high potential for runoff interception by upslope neighbours exerted strong detrimental effects on the water and phosphorus status of downslope S. tenacissima tussocks. Foliar delta(15)N values became more enriched with increasing soil depth. Multiple stepwise regression showed that competition potential and/or rhizosphere soil depth accounted for large proportions of variance in foliar delta(13)C, delta(18)O and delta(15)N among target tussocks (57, 37 and 64%, respectively). The results presented here highlight the key role that spatial redistribution of resources (water and nutrients) by runoff plays in semi-arid ecosystems. It is concluded that combined measurement of delta(13)C, delta(18)O and nutrient concentrations in bulk leaf tissue can provide insight into the intensity of competitive interactions occurring in natural plant communities.

Global patterns of foliar nitrogen isotopes and their relationships with climate, mycorrhizal fungi, foliar nutrient concentrations, and nitrogen availability

Ratios of nitrogen (N) isotopes in leaves could elucidate underlying patterns of N cycling across ecological gradients. To better understand global-scale patterns of N cycling, we compiled data on foliar N isotope ratios (delta(15)N), foliar N concentrations, mycorrhizal type and climate for over 11,000 plants worldwide. Arbuscular mycorrhizal, ectomycorrhizal, and ericoid mycorrhizal plants were depleted in foliar delta(15)N by 2 per thousand, 3.2 per thousand, 5.9 per thousand, respectively, relative to nonmycorrhizal plants. Foliar delta(15)N increased with decreasing mean annual precipitation and with increasing mean annual temperature (MAT) across sites with MAT >or= -0.5 degrees C, but was invariant with MAT across sites with MAT < -0.5 degrees C. In independent landscape-level to regional-level studies, foliar delta(15)N increased with increasing N availability; at the global scale, foliar delta(15)N increased with increasing foliar N concentrations and decreasing foliar phosphorus (P) concentrations. Together, these results suggest that warm, dry ecosystems have the highest N availability, while plants with high N concentrations, on average, occupy sites with higher N availability than plants with low N concentrations. Global-scale comparisons of other components of the N cycle are still required for better mechanistic understanding of the determinants of variation in foliar delta(15)N and ultimately global patterns in N cycling.

An assessment of the relationship between potential chemical indices of nitrogen saturation and nitrogen deposition in hardwood forests in southern Ontario

Southern Ontario receives the highest levels of atmospheric nitrogen (N) deposition in Canada and there are concerns that forests in the region may be approaching a state of 'N saturation'. In order to evaluate whether potential chemical indices provide evidence of N saturation, 23 hardwood plots were sampled along a modeled N-deposition gradient ranging from 9.3 to 12.8 kg/ha/year. All plots were dominated by sugar maple (Acer saccharum Marsh.) and foliar N and foliar delta(15)N were positively correlated with modeled N deposition. However, forest floor N content and the C:N ratio were unrelated to N deposition, but were instead related to soil pH and annual temperature; lower C:N ratios and higher N content in the forest floor were found at the most acidic sites in the cooler, northern part of the study region despite lower N deposition. Likewise, delta(15)N values in surface mineral soil and the (15)N enrichment factor of foliage (delta(15)N foliage - delta(15)N soil) are correlated to soil pH and temperature and not N deposition. Further, potential N mineralization, ammonification, and nitrification in Ontario maple stands were highest in the northern part of the region with the lowest modeled N deposition. Nitrogen cycling in soil appears to be primarily influenced by the N status of the forest floor and other soil properties rather than N deposition, indicating that chemical indices in soil in these hardwood plots may not provide an early indicator of N saturation.

Nitrogen fixation by Elaeagnus angustifolia in the reclamation of degraded croplands of Central Asia

Extensive degradation of irrigated croplands, due to increasing soil salinity and depletion of soil nutrient stocks, is a major problem in Central Asia (CA), one of the largest irrigated areas in the world. To assess the potential for improving the productive capacity of degraded lands by afforestation, we examined N(2) fixation of Elaeagnus angustifolia L. in mixed plantations with non-fixing Populus euphratica Oliv. and Ulmus pumila L. Fixation of N(2) was quantified by the (15)N natural abundance technique based on both foliar and whole-plant sampling during five consecutive growing seasons. Despite elevated root-zone soil salinity (6-10 dS m(-1)) and deficiency in plant-available P (4-15 mg kg(-1)), N(2) fixation (%Ndfa) increased from an initial value of 20% to almost 100% over 5 years. Within each growing season, %Ndfa steadily increased and peaked in the fall. Annual N(2) fixation, determined using foliar delta(15)N, initially averaged 0.02 Mg ha(-1), peaked at 0.5 Mg ha(-1) during the next 2 years and thereafter stabilized at 0.3 Mg ha(-1). Estimates based on whole-plant delta(15)N were <10% lower than those based on foliar delta(15)N. The increase in plant-available soil N was significantly higher in E. angustifolia plots than in P. euphratica and U. pumila plots. Increases in the concentrations of organic C (19%), total N (21%) and plant-available P (74%) in the soil were significant irrespective of tree species. This improvement in soil fertility is further evidence that afforestation with mixed-species plantations can be a sustainable land use option for the degraded irrigated croplands in CA.

Coordination of leaf structure and gas exchange along a height gradient in a tall conifer

The gravitational component of water potential and frictional resistance during transpiration lead to substantial reductions in leaf water potential (Psi(l)) near the tops of tall trees, which can influence both leaf growth and physiology. We examined the relationships between morphological features and gas exchange in foliage collected near the tops of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees of different height classes ranging from 5 to 55 m. This sampling allowed us to investigate the effects of tree height on leaf structural characteristics in the absence of potentially confounding factors such as irradiance, temperature, relative humidity and branch length. The use of cut foliage for measurement of intrinsic gas-exchange characteristics allowed identification of height-related trends without the immediate influences of path length and gravity. Stomatal density, needle length, needle width and needle area declined with increasing tree height by 0.70 mm(-2) m(-1), 0.20 mm m(-1), 5.9 x 10(-3) mm m(-1) and 0.012 mm(2) m(-1), respectively. Needle thickness and mesophyll thickness increased with tree height by 4.8 x 10(-2) mm m(-1) and 0.74 microm m(-1), respectively. Mesophyll conductance (g(m)) and CO(2) assimilation in ambient [CO(2)] (A(amb)) decreased by 1.1 mmol m(-2) s(-1) per m and 0.082 micromol m(-2) s(-1) per m increase in height, respectively. Mean reductions in g(m) and A(amb) of foliage from 5 to 55 m were 47% and 42%, respectively. The observed trend in A(amb) was associated with g(m) and several leaf anatomic characteristics that are likely to be determined by the prevailing vertical tension gradient during foliar development. A linear increase in foliar delta(13)C values with height (0.042 per thousand m(-1)) implied that relative stomatal and mesophyll limitations of photosynthesis in intact shoots increased with height. These data suggest that increasing height leads to both fixed structural constraints on leaf gas exchange and dynamic constraints related to prevailing stomatal behavior.

Analyses of 13C and 18O in tree rings of Callitris columellaris provide evidence of a change in stomatal control of photosynthesis in response to regional changes in climate

We examined relationships between stable isotopes of carbon (delta(13)C) and oxygen (delta(18)O) in tree rings of Callitris columellaris F. Muell in the semi-arid Pilbara region of north-western Australia. To test the hypothesis that stomatal control of photosynthesis decreases during drier periods, we developed delta(13)C and delta(18)O chronologies spanning 1919-1999, and used a permutation regression approach to relate a 21-year running correlation between delta(13)C and delta(18)O to rainfall and temperature at Marble Bar and our study site. The relationship between delta(13)C and delta(18)O switched from being always negative before 1955 to being consistently positive after 1976, suggesting an increase in stomatal control of photosynthesis in recent decades. Changes in the delta(13)C-delta(18)O relationship reflected changes in rainfall, which has increased in the region by 30% since 1976. The correlation between delta(13)C and delta(18)O was positively related to the 21-year running mean of normalized rainfall anomalies at both the study site (P = 0.045, Adj. r(2) = 0.47) and Marble Bar (P = 0.046, Adj. r(2) = 0.48). In addition, the delta(13)C-delta(18)O correlation was negatively related (P = 0.047, Adj. r(2) = 0.61) to temperatures at Marble Bar. Our interpretation of the role of changes in climate affecting the relationship between tree-ring delta(13)C and delta(18)O is supported by evidence from the isotope composition of foliage samples: foliar delta(13)C and delta(18)O were negatively correlated with log stomatal conductance (delta(13)C, r = -0.41; delta(18)O, r = -0.42), whereas the correlation between foliar delta(13)C and delta(18)O was positive (r = 0.63, P = 0.027) after the summer wet period. Our data indicate that stomatal control of photosynthesis in Callitris adjusts to region-wide changes in climate and that, in a warmer and drier world, trees might adapt by increasing non-stomatal control of photosynthesis.

Foliar 13C and 18O reveal differential physiological responses of canopy foliage to pre-planting weed control in a young spotted gum (Corymbia citriodora subsp. Variegata) plantation

Weed control may improve the growth of forest plantations by influencing soil water and nutrient availability, but our knowledge of leaf-level physiological responses to weed control at different within-canopy positions is limited for tropical and subtropical plantations. Foliar carbon (delta(13)C) and oxygen (delta(18)O) isotope compositions, gas exchange, and nitrogen (N(mass)) and phosphorus (P(mass)) concentrations at four canopy positions were assessed in a young spotted gum (Corymbia citriodora subsp. Variegata (F. Muell.) A.R. Bean & M.W. McDonald) plantation subjected to either weed control or no weed control treatment, to test if leaves at different positions within the tree canopy had the same physiological responses to the weed control treatment. Weed control increased foliar delta(13)C but lowered delta(18)O in the upper-outer and upper-inner canopy, indicating that weed control resulted in a higher foliar photosynthetic capacity at upper-canopy positions, a conclusion confirmed by gas exchange measurements. The increased photosynthetic capacity resulting from weed control can be explained by an increase in foliar N(mass). In the lower-outer canopy, weed control reduced foliar delta(13)C while lowering delta(18)O even more than in the upper-canopy, suggesting strong enhancement of the partial pressure of CO(2) in the leaf intercellular spaces and of foliar stomatal conductance in lower-canopy foliage. This conclusion was supported by gas exchange measurements. Foliar photosynthesis in the lower-inner canopy showed no significant response to weed control. The finding that leaves at different canopy positions differ in their physiological responses to weed control highlights the need to consider the canopy position effect when examining competition for soil nutrient and water resources between weeds and trees.

Foliar delta(15)N values characterize soil N cycling and reflect nitrate or ammonium preference of plants along a temperate grassland gradient.

The natural abundance of stable 15N isotopes in soils and plants is potentially a simple tool to assess ecosystem N dynamics. Several open questions remain, however, in particular regarding the mechanisms driving the variability of foliar δ15N values of non-N2 fixing plants within and across ecosystems. The goal of the work presented here was therefore to: (1) characterize the relationship between soil net mineralization and variability of foliar Δδ15N (δ15Nleaf − δ15Nsoil) values from 20 different plant species within and across 18 grassland sites; (2) to determine in situ if a plant’s preference for NO3− or NH4+ uptake explains variability in foliar Δδ15N among different plant species within an ecosystem; and (3) test if variability in foliar Δδ15N among species or functional group is consistent across 18 grassland sites. Δδ15N values of the 20 different plant species were positively related to soil net mineralization rates across the 18 sites. We found that within a site, foliar Δδ15N values increased with the species’ NO3− to NH4+ uptake ratios. Interestingly, the slope of this relationship differed in direction from previously published studies. Finally, the variability in foliar Δδ15N values among species was not consistent across 18 grassland sites but was significantly influenced by N mineralization rates and the abundance of a particular species in a site. Our findings improve the mechanistic understanding of the commonly observed variability in foliar Δδ15N among different plant species. In particular we were able to show that within a site, foliar δ15N values nicely reflect a plant’s N source but that the direction of the relationship between NO3− to NH4+ uptake and foliar Δδ15N values is not universal. Using a large set of data, our study highlights that foliar Δδ15N values are valuable tools to assess plant N uptake patterns and to characterize the soil N cycle across different ecosystems.

Quantifying foliar uptake of gaseous nitrogen dioxide using enriched foliar δ15N values

The magnitude and impact of gaseous nitrogen dioxide (NO(2)) directly entering the leaves were investigated using foliar nitrogen isotopic composition (delta(15)N) values in tomato (Lycopersicon esculentum) and tobacco (Nicotiana tabacum). Using a hydroponics-fumigation system, (15)NO(2) (20 and 40 ppb) was supplied to shoot systems and (50 and 500 microM) was supplied to root systems. Morphological, stable isotope and nitrate reductase activity (NRA) analyses were used to quantify foliar NO(2) uptake and to examine whether realistic concentrations of NO(2) influenced plant metabolism. Nicotiana tabacum and L. esculentum incorporated 15 and 11%, respectively, of (15)NO(2)-N into total biomass via foliar uptake under low supply. On a mass basis, N. tabacum and L. esculentum incorporated 3.3 +/- 0.9 and 3.1 +/- 0.8 mg of (15)NO(2)-N into biomass, respectively, regardless of availability. There were no strong effects on biomass accumulation or allocation, leaf delta(13)C values, or leaf or root NRA in response to NO(2) exposure. Foliar NO(2 )uptake may contribute a significant proportion of N to plant metabolism under N-limited conditions, does not strongly influence growth at 40 ppb, and may be traced using foliar delta(15)N values.

DRAINAGE AFFECTS TREE GROWTH AND C AND N DYNAMICS IN A MINEROTROPHIC PEATLAND

The lowering of the water table resulting from peatland drainage may dramatically alter C and N cycling in peatland ecosystems, which contain one-third of the total terrestrial C. In this study, tree annual ring width and C (delta(13)C) and N (delta(15)N) isotope ratios in soil and plant tissues (tree foliage, growth rings, and understory foliage) in a black spruce-tamarack (Picea mariana-Larix laricina) mixed-wood forest were examined to study the effects of drainage on tree growth and C and N dynamics in a minerotrophic peatland in west-central Alberta, Canada. Drainage increased the delta(15)N of soil NH4+ from a range of +0.6% per hundred to +2.9% per hundred to a range of +4.6% per hundred to +7.0% per hundred most likely through increased nitrification following enhanced mineralization. Plant uptake of 15N-enriched NH4+ in the drained treatment resulted in higher plant delta15N (+0.8% per hundred to +1.8% per hundred in the drained plots and -3.9% per hundred to -5.4% per hundred in the undrained plots), and deposition of litterfall N enriched with 15N increased the delta15N of total soil N in the surface layer in the drained (+2.9% per hundred) as compared with that in the undrained plots (+0.6% per hundred). The effect of drainage on foliar delta(13)C was species-specific, i.e., only tamarack showed a considerably less negative foliar delta(13)C in the drained (-28.1% per hundred) than in the undrained plots (-29.1% per hundred), indicating improved water use efficiency (WUE) by drainage. Tree ring area increments were significantly increased following drainage, and delta(13)C and delta(15)N in tree growth rings of both species showed responses to drainage retrospectively. Tree-ring delta(13)C data suggested that drainage improved WUE of both species, with a greater and more prolonged response in tamarack than in black spruce. Our results indicate that drainage caused the studied minerotrophic peatland to become a more open ecosystem in terms of C and N cycling and loss. The effects of forested peatland drainage or drying on C and N balances deserve further research in order to better understand their roles in future global change.

Water source partitioning among trees growing on shallow karst soils in a seasonally dry tropical climate

The sources of water used by woody vegetation growing on karst soils in seasonally dry tropical regions are little known. In northern Yucatan (Mexico), trees withstand 4-6 months of annual drought in spite of the small water storage capacity of the shallow karst soil. We hypothesized that adult evergreen trees in Yucatan tap the aquifer for a reliable supply of water during the prolonged dry season. The naturally occurring concentration gradients in oxygen and hydrogen stable isotopes in soil, bedrock, groundwater and plant stem water were used to determine the sources of water used by native evergreen and drought-deciduous tree species. While the trees studied grew over a permanent water table (9-20 m depth), pit excavation showed that roots were largely restricted to the upper 2 m of the soil/bedrock profile. At the peak of the dry season, the delta(18)O signatures of potential water sources for the vegetation ranged from 4.1 +/- 1.1 per thousand in topsoil to -4.3 +/- 0.1 per thousand in groundwater. The delta(18)O values of tree stem water ranged from -2.8 +/- 0.3 per thousand in Talisia olivaeformis to 0.8 +/- 1 per thousand in Ficus cotinifolia, demonstrating vertical partitioning of soil/bedrock water among tree species. Stem water delta(18)O values were significantly different from that of groundwater for all the tree species investigated. Stem water samples plotted to the right of the meteoric water line, indicating utilization of water sources subject to evaporative isotopic enrichment. Foliar delta(13)C in adult trees varied widely among species, ranging from -25.3 +/- 0.3 per thousand in Enterolobium cyclocarpum to -28.7 +/- 0.4 per thousand in T. olivaeformis. Contrary to initial expectations, data indicate that native trees growing on shallow karst soils in northern Yucatan use little or no groundwater and depend mostly on water stored within the upper 2-3 m of the soil/bedrock profile. Water storage in subsurface soil-filled cavities and in the porous limestone bedrock is apparently sufficient to sustain adult evergreen trees throughout the pronounced dry season.

Hydraulic architecture and photosynthetic capacity as constraints on release from suppression in Douglas-fir and western hemlock

We compared hydraulic architecture, photosynthesis and growth in Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), a shade-intolerant species, and western hemlock (Tsuga heterophylla (Raf.) Sarg.), a shade-tolerant species, to study the temporal pattern of release from suppressive shade. In particular, we sought to determine whether hydraulic architecture or photosynthetic capacity is most important in constraining release. The study was conducted at two sites with mixed stands of 10- to 20-year-old Douglas-fir and western hemlock. At one site, the stand had been thinned allowing release of the understory trees, whereas at the other site, the stand remained unthinned. Douglas-fir had lower height growth (from 1998-2003) and lower relative height growth (height growth from 1998 to 2003/height in 1998) than western hemlock. However, relative height growth of released versus suppressed trees was higher in Douglas-fir (130%) than in western hemlock (65%), indicating that, although absolute height growth was less, Douglas-fir did release from suppression. Release seemed to be constrained initially by a limited photosynthetic capacity in both species. Five years after release, Douglas-fir trees had 14 times the leaf area and 1.5 times the leaf nitrogen concentration (N (area)) of suppressed trees. Needles of released western hemlock trees had about twice the maximum assimilation rate (A (max)) at ambient [CO(2)] as needles of suppressed trees and exhibited no photoinhibition at the highest irradiances. After release, trees increased in leaf area, leaf N concentration and overall photosynthetic capacity. Subsequently, hydraulic architecture appeared to constrain release in Douglas-fir and, to a lesser extent, in western hemlock. Released trees had significantly less negative foliar delta(13)C values than suppressed trees and showed a positive relationship between leaf area:sapwood area ratio (A (L)/A (S)) and delta(13)C, suggesting that trees with more leaf area for a given sapwood area experienced a stomatal limitation on carbon gain. Nonetheless, these changes had no significant effects on leaf specific conductivities of suppressed versus released trees of either species, but leaf specific root conductance was significantly lower in released Douglas-fir.

Partitioning of water and nitrogen in co-occurring Mediterranean woody shrub species of different evolutionary history.

We studied the interspecific and intraspecific variation in the development of water stress and in the use of different water and nitrogen sources during the spring (wet season) and summer (dry season) in a shrub community in NE Spain. We measured shoot water potentials, stable deuterium isotopic composition (delta D) of xylem sap, leaf mass per area, leaf N and C concentrations, gas exchange, leaf delta(13)C, and leaf delta(15)N of the dominant species (Quercus coccifera, Arbutus unedo, Pistacia lentiscus, Erica multiflora, Globularia alypum). The delta D, the delta(13)C and the shoot water potential values showed diurnal, seasonal, intraspecific and interspecific variation in the source and use of water. There was also seasonal, intraspecific and interspecific variation in the foliar delta(15)N and N concentrations. In summer, some species (A. unedo, P. lentiscus and E. multiflora) presented significantly different delta D values in morning and afternoon measurements likely indicating that they used different sources of water during the day, and a dual root system in these species. We conjecture that dew may be one of these water sources. Species predawn water potential was negatively correlated with species xylem water delta D. There was also a positive correlation between delta(13)C and delta D in P. lentiscus, species for which we took additional samples from nearby sites. These results suggest that the access to water from greater depths allowed the maintenance of more favourable plant water supply. Multivariate principal component analysis based on the studied hydrological and isotope variables clearly separated the seasons (wet spring and dry summer) and the species. The species resulted separated according to their evolutionary history (Pre-Mediterranean and Mediterranean) and the associated root and functional traits. These results show water (and nitrogen) partitioning among coexisting species of the same functional type (Mediterranean woody shrubs). They also show the great intraspecific plasticity of responses to resource availability.

Delta15N values of tropical savanna and monsoon forest species reflect root specialisations and soil nitrogen status.

A large number of herbaceous and woody plants from tropical woodland, savanna, and monsoon forest were analysed to determine the impact of environmental factors (nutrient and water availability, fire) and biological factors (microbial associations, systematics) on plant delta(15)N values. Foliar delta(15)N values of herbaceous and woody species were not related to growth form or phenology, but a strong relationship existed between mycorrhizal status and plant delta(15)N. In woodland and savanna, woody species with ectomycorrhizal (ECM) associations and putative N(2)-fixing species with ECM/arbuscular (AM) associations had lowest foliar delta(15)N values (1.0-0.6 per thousand ), AM species had mostly intermediate delta(15)N values (average +0.6 per thousand ), while non-mycorrhizal Proteaceae had highest delta(15)N values (+2.9 to +4.1 per thousand ). Similar differences in foliar delta(15)N were observed between AM (average 0.1 and 0.2 per thousand ) and non-mycorrhizal (average +0.8 and +0.3 per thousand ) herbaceous species in woodland and savanna. Leguminous savanna species had significantly higher leaf N contents (1.8-2.5% N) than non-fixing species (0.9-1.2% N) indicating substantial N acquisition via N(2) fixation. Monsoon forest species had similar leaf N contents (average 2.4% N) and positive delta(15)N values (+0.9 to +2.4 per thousand ). Soil nitrification and plant NO(3)(-) use was substantially higher in monsoon forest than in woodland or savanna. In the studied communities, higher soil N content and nitrification rates were associated with more positive soil delta(15)N and plant delta(15)N. In support of this notion, Ficus, a high NO(3)(-) using taxa associated with NO(3)(-) rich sites in the savanna, had the highest delta(15)N values of all AM species in the savanna. delta(15)N of xylem sap was examined as a tool for studying plant delta(15)N relations. delta(15)N of xylem sap varied seasonally and between differently aged Acacia and other savanna species. Plants from annually burnt savanna had significantly higher delta(15)N values compared to plants from less frequently burnt savanna, suggesting that foliar (15)N natural abundance could be used as marker for assessing historic fire regimes. Australian woodland and savanna species had low leaf delta(15)N and N content compared to species from equivalent African communities indicating that Australian biota are the more N depauperate. The largest differences in leaf delta(15)N occurred between the dominant ECM Australian and African savanna (miombo) species, which were depleted and enriched in (15)N, respectively. While the depleted delta(15)N of Australian ECM species are similar to those of previous reports on ECM species in natural plant communities, the (15)N-enriched delta(15)N of African ECM species represent an anomaly.