Leaf Delta Research Articles

Latitudinal and climate effects on key plant traits in Chinese forest ecosystems

Abstracts A long-standing goal of ecology and forest management is to understand the environmental controls of resource utilization strategy of plants along geographical gradients. However, the mechanisms that guide this phenomenon, especially along latitude gradients, remain unclear. Using key leaf functional trait data of 1185 species within 120 sites in Eastern and Western China, we quantified the relationships between plant traits, namely leaf area (LA), specific leaf area (SLA), leaf nitrogen content, leaf phosphorus content, leaf dry matter content (LDMC) and leaf delta 13C (d 13C) content, and local climate along the latitude in Eastern and Western China. Our results revealed that: (1) Plant traits showed a wide variation across the study locations in China. The variation in plant traits was strongly correlated with latitude. (2) In Eastern China, latitude alone explained 54% and 53% of the variation in N:P and leaf phosphorus content (Pmass (g/kg)), respectively. In Western China, latitude alone explained 75% of the variation in leaf area. In Eastern China, most plant traits are significantly correlated with each other. However, this phenomenon is not obvious in Western China. (3) Immediate climate factors (mean climate in the month when the leaf traits were sampled) contributes most to explaining functional traits related to leaf nutrients, such as leaf nitrogen content (Nmass (g/kg), Narea (g/cm2)), leaf phosphorus content (Pmass (g/kg), Parea (g/cm2)), and annual mean climate contributes more to explaining functional traits related to leaf survival strategies. (4) In Eastern China, immediate climate and annual mean climate variables can explain the variance of 39.2% and 41.3% of plant traits variation, respectively. In Western China, the effect size of corresponding factors are 23.4% and 25.2%, respectively.

Patterns in leaf traits of leguminous and non-leguminous dominant trees along a rainfall gradient in Ghana

Aims Both dominance distribution of species and the composition of the dominant species determine the distribution of traits within community. Leaf carbon (C) and nitrogen (N) isotopic composition are important leaf traits, and such traits of dominant species are associated with ecosystem C, water and N cycling. Very little is known how dominant species with distinct traits (e.g. N-fixing leguminous and non-leguminous trees) mediate resource utilization of the ecosystems in stressful environment. Methods Leaves of 81 dominant leguminous and non-leguminous trees were collected in forest (moist semi-deciduous and dry semi-deciduous ecosystems) and savanna (costal savanna, Guinean savanna and west Sudanian savanna ecosystems) areas and the transitional zone (between the forest and the savanna) along the transect from the south to the north of Ghana. We measured leaf traits, i.e. leaf delta C-13, leaf delta N-15, leaf water content, leaf mass per area (LMA) and C and N concentration. Correlation analyses were used to examine trait-trait relationships, and relationships of leaf traits with temperature and precipitation. We used analysis of covariance to test the differences in slopes of the linear regressions between legumes and non-legumes. Important Findings Leaf delta C-13, delta N-15, leaf water content and LMA did not differ between leguminous and non-leguminous trees. Leaf N concentration and C: N ratio differed between the two groups. Moreover, leaf traits varied significantly among the six ecosystems. delta C-13 values were negatively correlated with annual precipitation and positively correlated with mean annual temperature. In contrast, leaf delta N-15 of non-leguminous trees were positively correlated with annual precipitation and negatively correlated with mean annual temperature. For leguminous trees, such correlations were not significant. We also found significant coordination between leaf traits. However, the slopes of the linear relationships were significantly different between leguminous and non-leguminous trees. Our results indicate that shifts in dominant trees with distinct water-use efficiency were corresponded to the rainfall gradient. Moreover, leguminous trees, those characterized with relative high water-use efficiency in the low rainfall ecosystems, were also corresponded to the relative high N use efficiency. The high proportion of leguminous trees in the savannas is crucial to mitigate nutrient stress.

Ants mediate foliar structure and nitrogen acquisition in a tank‐bromeliad

Aechmea mertensii is a tank-bromeliad that roots on ant-gardens initiated by the ants Camponotus femoratus and Pachycondyla goeldii. Its leaves form compartments acting as phytotelmata that hold rainwater and provide habitats for invertebrates. In this article, we aimed to determine whether the association with either C. femoratus or P. goeldii influenced the vegetative traits of A. mertensii, invertebrate diversity and nutrient assimilation by the leaves. Transmitted light, vegetative traits and phytotelmata contents were compared between the two A. mertensii ant-gardens. Camponotus femoratus colonized partially shaded areas, whereas P. goeldii colonized exposed areas. The bromeliads' rosettes had a large canopy (C. femoratus ant-gardens), or were smaller and amphora shaped (P. goeldii ant-gardens). There were significant differences in leaf anatomy, as shaded leaves were thicker than exposed leaves. The mean volumes of water, fine particulate organic matter and detritus in C. femoratus-associated bromeliads were three to five times higher than in P. goeldii-associated bromeliads. Moreover, the highest invertebrate diversity and leaf delta(15)N values were found in C. femoratus-associated bromeliads. This study enhances our understanding of the dynamics of biodiversity, and shows how ant-plant interactions can have trophic consequences and thus influence the architecture of the interacting plant via a complex feedback loop.

Greater impact of extreme drought on photosynthesis of grasslands exposed to a warmer climate in spite of acclimation

In view of the projected increase in the frequency of extreme events during this century, we investigated the impact of a drought extreme on leaf ecophysiological parameters and carbon isotope composition (delta(13)C) of grassland communities with species richness (S) of one, three or nine species. The communities, grown for 3 years at either ambient air temperatures (ambient T(air)) or ambient T(air) + 3 degrees C (elevated T(air)), were additionally subjected to an imposed drought by withholding water for 24 days. During the previous 3 years equal precipitation was applied in both temperature treatments, thus communities at elevated T(air) had experienced more frequent, mild droughts. However, it was unknown whether this resulted in a higher resistance for facing extreme droughts. At similar soil matric potentials stomatal conductance (g(s)) and transpiration (Tr) were higher at elevated than ambient T(air), indicating acclimation to lower soil water content. Despite the stomatal acclimation observed, plants in elevated T(air) showed a lower resistance to the drought extreme as indicated by their lower photosynthetic rate (A(max)), g(s) and Tr during the entire duration of the drought extreme. Lower values for A(max), Tr and g(s) were also recorded in species at S = 3 as compared with species at S = 1 for both temperature treatments, but no further differences with S = 9 suggesting that stress was not alleviated at higher S-levels. The discrimination of (13)C was poorly correlated with measurements of instantaneous leaf water-use efficiency (A(max)/Tr) and, with this time scale and sampling method, it was not possible to detect any potential change in plant water-use efficiency using leaf delta(13)C.

Leaf δ13C reflects ecosystem patterns and responses of alpine plants to the environments on the Tibetan Plateau

Leaf δ13C is an indicator of water‐use efficiency and provides useful information on the carbon and water balance of plants over longer periods. Variation in leaf δ13C between or within species is determined by plant physiological characteristics and environmental factors. We hypothesized that variation in leaf δ13C values among dominant species reflected ecosystem patterns controlled by large‐scale environmental gradients, and that within‐species variation indicates plant adaptability to environmental conditions. To test these hypotheses, we collected leaves of dominant species from six ecosystems across a horizontal vegetation transect on the Tibetan Plateau, as well as leaves of Kobresia pygmaea (herbaceous) throughout its distribution and leaves of two coniferous tree species (Picea crassifolia, Abies fabri) along an elevation gradient throughout their distribution in the Qilian Mountains and Gongga Mountains, respectively. Leaf δ13C of dominant species in the six ecosystems differed significantly, with values for evergreen coniferous<evergreen broadleaved tree<alpine shrub<sedges∼graminoid<xeromorphs. Leaf δ13C values of the dominant species and of K. pygmaea were negatively correlated with annual precipitation along a water gradient, but leaf δ13C of A. fabri was not significantly correlated with precipitation in habitats without water‐stress. This confirms that variation of δ13C between or within species reflects plant responses to environmental conditions. Leaf δ13C of the dominant species also reflected water patterns on the Tibetan Plateau, providing evidence that precipitation plays a primary role in controlling ecosystem changes from southeast to northwest on the Tibetan Plateau.

Impact of epidermal leaf mining by the aspen leaf miner (Phyllocnistis populiella) on the growth, physiology, and leaf longevity of quaking aspen

The aspen leaf miner, Phyllocnistis populiella, feeds on the contents of epidermal cells on both top (adaxial) and bottom (abaxial) surfaces of quaking aspen leaves, leaving the photosynthetic tissue of the mesophyll intact. This type of feeding is taxonomically restricted to a small subset of leaf mining insects but can cause widespread plant damage during outbreaks. We studied the effect of epidermal mining on aspen growth and physiology during an outbreak of P. populiella in the boreal forest of interior Alaska. Experimental reduction of leaf miner density across two sites and 3 years significantly increased annual aspen growth rates relative to naturally mined controls. Leaf mining damage was negatively related to leaf longevity. Leaves with heavy mining damage abscised 4 weeks earlier, on average, than leaves with minimal mining damage. Mining damage to the top and bottom surfaces of leaves had different effects on physiology. Mining on the top surface of the leaf had no significant effect on photosynthesis or conductance and was unrelated to leaf stable C isotope ratio (delta(13)C). Mining damage to the bottom leaf surface, where stomata are located, had significant negative effects on net photosynthesis and water vapor conductance. Percent bottom mining was positively related to leaf delta(13)C. Taken together, the data suggest that the primary mechanism for the reduction of photosynthesis by epidermal leaf mining by P. populiella is the failure of stomata to open normally on bottom-mined leaves.

C4 photosynthetic isotope exchange in NAD-ME- and NADP-ME-type grasses

Monitoring photosynthetic isotope exchange is an important tool for predicting the influence of plant communities on the global carbon cycle in response to climate change. C(4) grasses play an important role in the global carbon cycle, but their contribution to the isotopic composition of atmospheric CO(2) is not well understood. Instantaneous measurements of (13)CO(2) (Delta(13)C) and C(18)OO (Delta(18)O) isotope exchange in five NAD-ME and seven NADP-ME C(4) grasses have been conducted to investigate the difference in photosynthetic CO(2) isotopic fractionation in these subgroups. As previously reported, the isotope composition of the leaf material (delta(13)C) was depleted in (13)C in the NAD-ME compared with the NADP-ME grasses. However, Delta(13)C was not different between subtypes at high light, and, although Delta(13)C increased at low light, it did so similarly in both subtypes. This suggests that differences in leaf delta(13)C between the C(4) subtypes are not caused by photosynthetic isotope fractionation and leaf delta(13)C is not a good indicator of bundle sheath leakiness. Additionally, low carbonic anhydrase (CA) in C(4) grasses may influences Delta(13)C and should be considered when estimating the contribution of C(4) grasses to the global isotopic signature of atmospheric CO(2). It was found that measured Delta(18)O values were lower than those predicted from leaf CA activities and Delta(18)O was similar in all species measured. The Delta(18)O in these C(4) grasses is similar to low Delta(18)O previously measured in C(4) dicots which contain 2.5 times the leaf CA activity, suggesting that leaf CA activity is not a predictor of Delta(18)O in C(4) plants.

Variation in woody plant δ13C along a topoedaphic gradient in a subtropical savanna parkland

delta(13)C values of C(3) plants are indicators of plant carbon-water relations that integrate plant responses to environmental conditions. However, few studies have quantified spatial variation in plant delta(13)C at the landscape scale. We determined variation in leaf delta(13)C, leaf nitrogen per leaf area (N(area)), and specific leaf area (SLA) in April and August 2005 for all individuals of three common woody species within a 308 x 12-m belt transect spanning an upland-lowland topoedaphic gradient in a subtropical savanna in southern Texas. Clay content, available soil moisture, and soil total N were all negatively correlated with elevation. The delta(13)C values of Prosopis glandulosa (deciduous N(2)-fixing tree legume), Condalia hookeri (evergreen shrub), and Zanthoxylum fagara (evergreen shrub) leaves increased 1-4 per thousand with decreasing elevation, with the delta(13)C value of P. glandulosa leaves being 1-3 per thousand higher than those of the two shrub species. Contrary to theory and results from previous studies, delta(13)C values were highest where soil water was most available, suggesting that some other variable was overriding or interacting with water availability. Leaf N(area) was positively correlated with leaf delta(13)C of all species (p < 0.01) and appeared to exert the strongest control over delta(13)C along this topoedaphic gradient. Since leaf N(area) is positively related to photosynthetic capacity, plants with high leaf N(area) are likely to have low p (I)/p (a) ratios and therefore higher delta(13)C values, assuming stomatal conductance is constant. Specific leaf area was not correlated significantly with leaf delta(13)C. Following a progressive growing season drought in July/August, leaf delta(13)C decreased. The lower delta(13)C in August may reflect the accumulation of (13)C-depleted epicuticular leaf wax. We suggest control of leaf delta(13)C along this topoedaphic gradient is mediated by leaf N(area) rather than by stomatal conductance limitations associated with water availability.

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.

Roles of soil chemistry and water availability in site-related 13C variations in French beech forests

The carbon isotopic composition (delta(13)C) of wood and leaf cellulose of beech trees (Fagus sylvatica L.) was studied at 80 sites in northeastern France. We sampled sites with contrasting water balance, depending on soil type and precipitation. We tested the hypothesis that inter-site variations in plant delta(13)C reflect the spatial distribution of soil water availability, and we assessed whether delta(13)C could be used as a bioindicator of soil water availability. Patterns of variation in delta(13)C were compared with estimates of monthly water balance and with other soil characteristics. Between-site variability in delta(13)C was high (2.9 per thousand range in wood cellulose, 2.1 per thousand in leaf cellulose), but variation in water availability appeared to be only a minor factor contributing to this variation in delta(13)C. Unexpectedly, spatial variations in wood and leaf cellulose delta(13)C were significantly and positively related to soil fertility expressed by soil pH (r = 0.42 and 0.43, respectively) and cation content. On average, trees growing on acidic soils displayed 0.5 per thousand lower delta(13)C in both wood and leaf material than trees growing on neutral or calcareous soils. Our initial hypothesis of a strong negative relationship between delta(13)C and site water availability was not confirmed. In the study zone, neither wood nor leaf delta(13)C appeared to be a reliable bioindicator of spatial variations in water availability. Possible causes for the lack of a relationship are discussed. Our findings confirm, under natural conditions, the strong effect of soil fertility on water-use efficiency previously observed in experiments. This effect needs to be considered in isotopic studies involving different sites.

CONTRASTING PLANT PHYSIOLOGICAL ADAPTATION TO CLIMATE IN THE NATIVE AND INTRODUCED RANGE OFHYPERICUM PERFORATUM

How introduced plants, which may be locally adapted to specific climatic conditions in their native range, cope with the new abiotic conditions that they encounter as exotics is not well understood. In particular, it is unclear what role plasticity versus adaptive evolution plays in enabling exotics to persist under new environmental circumstances in the introduced range. We determined the extent to which native and introduced populations of St. John's Wort (Hypericum perforatum) are genetically differentiated with respect to leaf-level morphological and physiological traits that allow plants to tolerate different climatic conditions. In common gardens in Washington and Spain, and in a greenhouse, we examined clinal variation in percent leaf nitrogen and carbon, leaf delta(13)C values (as an integrative measure of water use efficiency), specific leaf area (SLA), root and shoot biomass, root/shoot ratio, total leaf area, and leaf area ratio (LAR). As well, we determined whether native European H. perforatum experienced directional selection on leaf-level traits in the introduced range and we compared, across gardens, levels of plasticity in these traits. In field gardens in both Washington and Spain, native populations formed latitudinal clines in percent leaf N. In the greenhouse, native populations formed latitudinal clines in root and shoot biomass and total leaf area, and in the Washington garden only, native populations also exhibited latitudinal clines in percent leaf C and leaf delta(13)C. Traits that failed to show consistent latitudinal clines instead exhibited significant phenotypic plasticity. Introduced St. John's Wort populations also formed significant or marginally significant latitudinal clines in percent leaf N in Washington and Spain, percent leaf C in Washington, and in root biomass and total leaf area in the greenhouse. In the Washington common garden, there was strong directional selection among European populations for higher percent leaf N and leaf delta(13)C, but no selection on any other measured trait. The presence of convergent, genetically based latitudinal clines between native and introduced H. perforatum, together with previously published molecular data, suggest that native and exotic genotypes have independently adapted to a broad-scale variation in climate that varies with latitude.

Does the Prostrate-leaved Geophyte Brunsvigia orientalis Utilize Soil-derived CO2 for Photosynthesis?

A test was made of the hypothesis that the prostrate growth habit of the leaves of the geophyte Brunsvigia orientalis enables utilization of soil-derived CO(2) and is related to the presence of lysigenous air-filled channels characteristic of B. orientalis leaves. Brunsvigia orientalis was sampled at a field site. Leaf anatomy, stomatal density, leaf/soil gas exchange characteristics and soil atmosphere and leaf delta(13)C isotope abundances were examined. The leaves of B. orientalis have large lysigenous air-filled channels separating the upper and lower surfaces of the leaves. The upper surface comprised approx. 70 % of the leaf mass and 75 % of the leaf N (mmol g(-1)). Between 20 % and 30 % of the stomatal conductance and CO(2) assimilation was through the lower surface of the leaf. CO(2) efflux rates from the soil surface were up to 5.4 micromol m(-2) s(-1) while photosynthetic fluxes through the lower surface of the leaves were approx. 7 micromol m(-2) s(-1). However, the utilization of soil-derived CO(2) only altered the leaf delta(13)C isotope abundance of the prostrate leaves by a small amount. Using delta(13)C values it was estimated that 7 % of the leaf tissue C was derived from soil-derived CO(2). A small proportion of photosynthetically fixed CO(2) was derived from the soil, with minimal associated transpirational H(2)O loss into the space between the leaf and soil. The soil-derived CO(2), taken up through the lower surface was probably assimilated by the palisade tissue in the upper surface of the leaf which was exposed to sunlight and where most of the leaf N was located. The occurrence of lysigenous air channels in the leaves may provide longitudinal strength without impaired transfer of CO(2) taken up through the lower surface to the upper surface.

Variation in leaf physiology of Salix arctica within and across ecosystems in the High Arctic: test of a dual isotope (Δ13C and Δ18O) conceptual model

Leaf carbon isotope discrimination (Delta(13)C) varies with the balance between net photosynthesis (A) and stomatal conductance (g ( s )). Inferences that can be made with Delta(13)C are limited, as changes could reflect variation in A and/or g ( s ). Investigators have suggested that leaf delta(18)O enrichment above source water (Delta(18)O) may enable differentiation between sources of variation in Delta(13)C, as leaf Delta(18)O varies with transpiration rate (E), which is closely correlated with g ( s ) when leaves experience similar leaf to air vapor pressure differences. We examined leaf gas exchange of Salix arctica at eight sites with similar air temperatures and relative humidities but divergent soil temperatures and soil water contents near Pituffik, Greenland (76 degrees N, 38 degrees W). We found negative correlations at the site level between g ( s ) and Delta(18)O in bulk leaf tissue (r (2) = 0.62, slope = -17.9 per thousand/mol H(2)O m(-2) s(-1), P = 0.02) and leaf alpha-cellulose (r (2) = 0.83, slope = -11.5 per thousand mol H(2)O m(-2) s(-1), P < 0.01), consistent with the notion that leaf water enrichment declines with increasing E. We also found negative correlations at the site-level between intrinsic water-use efficiency (iWUE) and Delta(13)C in bulk leaf tissue (r (2) = 0.65, slope = -0.08 per thousand/mumol CO(2) /mol H(2)O, P = 0.02) and leaf alpha-cellulose (r (2) = 0.50, slope = -0.05 per thousand/[mumol CO(2) /mol H(2)O], P = 0.05). When increasing Delta(13)C was driven by increasing g ( s ) alone, we found negative slopes between Delta(13)C and Delta(18)O for bulk leaf tissue (-0.664) and leaf alpha-cellulose (-1.135). When both g ( s ) and A (max) increased, we found steeper negative slopes between Delta(13)C and Delta(18)O for bulk leaf tissue (-2.307) and leaf alpha-cellulose (-1.296). Our results suggest that the dual isotope approach is capable of revealing the qualitative contributions of g ( s ) and A (max) to Delta(13)C at the site level. In our study, bulk leaf tissue was a better medium than leaf alpha-cellulose for application of the dual isotope approach.

Genetic basis of phenotypic correlations among growth traits in hybrid willow (Salix dasyclados×S. viminalis) grown under two water regimes

Phenotypic correlations and quantitative trait loci (QTL) for important growth traits and a surrogate of intrinsic water-use efficiency (leaf delta(13)C) were analysed in a willow pedigree of 92 full-sibling clones grown under two water regimes. The major objective was to examine the genetic basis of the phenotypic correlations. Cuttings of Salix were glasshouse-grown during one growing season. The relative growth rate (RGR) and underlying traits were assessed. QTL analysis was conducted based on an available linkage map for Salix. Leaf area productivity and leaf nitrogen productivity were more important in determining RGR than leaf area ratio and specific leaf area. However, phenotypic correlations among growth traits partly varied between the two environments. QTL were detected for most growth traits, among them many common QTL for different traits. The QTL pattern reflected the phenotypic correlation pattern. None of the QTL for the complex traits was consistent across the different environments. The results demonstrate a genetic basis for phenotypic correlations among growth traits in Salix, and provide evidence for the existence of 'master switches' regulating some of the traits.

The effect of transient and continuous drought on yield, photosynthesis and carbon isotope discrimination in sugar beet (Beta vulgaris L.)

Stable carbon isotope discrimination (delta13C), photosynthetic performance (A), dry matter accumulation (DW), and sucrose yield (Y(s)) of sugar beet were evaluated in a glasshouse experiment under transient (TS) and permanent (PS) water stress. A was significantly reduced under drought, to an extent depending on stress duration. The reduced A was strictly associated with a low DW and Y(s), the later being 42% lower in PS than control plants (C). Restoring water steeply increased A and the associated leaf traits (RWC, leaf water potential etc.), but the increase of Y(s) was negligible. Therefore, the negative effects of severe water stress in the early growth period, though reversible on gas-exchange and most leaf traits, can drastically reduce Y(s) of sugar beet. Furthermore, A seems not to be effective in predicting sucrose accumulation, although it was very effective in detecting the occurrence of plant water stress. The A/C(i) model was used to assess the photosynthetic adjustments to continuous or transient drought by calculating the photosynthetic parameters Vcmax and Jmax and then compared with delta13C. Mesophyll conductance (g(m)) was estimated by comparing delta13C measured on soluble sugars and gas-exchange data. This approach confirmed the expectation that g(m) was limiting A and that there was a significant drop in [CO2] from the substomatal cavities and the chloroplast stroma both in favourable and drought conditions. Therefore, the carbon concentration at the carboxylation site was overestimated by 25-35% by conventional gas-exchange measurements, and Vcmax was consistently underestimated when g(m) was not taken into account, especially under severe drought. Root delta13C was found to be strictly related to sucrose content (brix%), Y(s) and root dry weight, and this was especially clear when delta13C was measured on bulk dry matter. By contrast, leaf delta13C measured in soluble sugars (delta(s)) and bulk dry matter (delta(dm)) were found to correlate weakly to brix% and yield, and this was not surprising as the integration time-scale of leaf delta(s) and delta(dm) were found to be shorter than that of root delta13C in bulk dry matter. The effect of water stress on diffusive and biochemical limitations with different integration times ranged from 1 d (leaf delta(s)) to more than 1 month (root delta(dm)).

Ecosystem implications of genetic variation in water-use of a dominant riparian tree.

Genetic variation in dominant species can affect plant and ecosystem functions in natural systems through multiple pathways. Our study focuses on how genetic variation in a dominant riparian tree ( Populus fremontii, P. angustifolia and their natural F(1) and backcross hybrids) affects whole-tree water use, and its potential ecosystem implications. Three major patterns were found. First, in a 12-year-old common garden with trees of known genetic makeup, hybrids had elevated daily integrated leaf-specific transpiration ( E(tl); P=0.013) and average canopy conductance ( G(c); P=0.037), with both E(tl) and G(c) approximately 30% higher in hybrid cross types than parental types. Second, delta(13)C values of leaves from these same trees were significantly more negative in hybrids ( P=0.004), and backcross hybrids had significantly more negative values than all other F(1) hybrid and parental types ( P<0.001). Third, in the wild, a similar pattern was found in leaf delta(13)C values where both hybrid cross types had the lowest values ( P<0.001) and backcross hybrids had lower delta(13)C values than any other tree type ( P<0.001). Our findings have two important implications: (1). the existence of a consistent genetic difference in whole-tree physiology suggests that whole-tree gas and water exchange could be another pathway through which genes could affect ecosystems; and (2). such studies are important because they seek to quantify the genetic variation that exists in basic physiological processes-such knowledge could ultimately place ecosystem studies within a genetic framework.

Stable carbon isotope composition of terrestrial leaves: inter‐ and intraspecies variability, cellulose and whole‐leaf tissue difference, and potential for climate reconstruction

AbstractStable carbon isotope analysis of terrestrial plant leaves preserved in Quaternary lake sediments has the potential to provide high‐resolution reconstructions of past climatic conditions. Preferably, δ13C measurements should be made on a single leaf component, e.g. cellulose, but this approach is often precluded by limited leaf availability. Previous work suggests that reliable palaeoclimatic information also may be derived from δ13C measurements on whole‐leaf tissue, given a similar degree of leaf decomposition between samples. Leaf δ13C data for 12 Scandinavian species of dwarf‐shrubs, shrubs and trees, and a comparison of δ13C data on recent and late Holocene Salix herbacea leaves, revealed that the δ13C signal registered by holocellulose is largely reflected by measurements on whole‐leaf tissue. Holocellulose was found to be consistently enriched in 13C, although this δ13C offset was smaller for subfossil leaves. This supports the use of δ13C measurements on whole‐leaf tissue for climate reconstruction, at least for leaves preserved in soft, late Holocene sediments with minimal diagenetic effects. Leaf carbon and nitrogen data on fresh leaves of the same 12 Scandinavian species, and corresponding data on late Holocene Salix herbacea leaves, suggest that the leaf C:N ratio is a suitable indicator of the degree of leaf decomposition. Copyright © 2003 John Wiley &amp; Sons, Ltd.

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.

N-15 natural abundance studies in Australian commercial sugarcane

The measurement of natural N-15 abundance is a well-established technique for the identification and quantification of biological N-2 fixation in plants. Associative N-2 fixing bacteria have been isolated from sugarcane and reported to contribute potentially significant amounts of N to plant growth and development. It has not been established whether Australian commercial sugarcane receives significant input from biological N-2 fixation, even though high populations of N-2 fixing bacteria have been isolated from Australian commercial sugarcane fields and plants. In this study, delta(15)N measurements were used as a primary measure to identify whether Australian commercial sugarcane was obtaining significant inputs of N via biological N-2 fixation. Quantification of N input, via biological N-2 fixation, was not possible since suitable non-N-2 fixing reference plants were not present in commercial cane fields. The survey of Australian commercially grown sugarcane crops showed the majority had positive leaf delta(15)N values (73% >3.00parts per thousand, 63% of which were <5.00parts per thousand), which was not indicative of biological N-2 fixation being the major source of N for these crops. However, a small number of sites had low or negative leaf delta(15)N values. These crops had received high N fertiliser applications in the weeks prior to sampling. Two possible pathways that could result in low delta(15)N values for sugarcane leaves (other than N-2 fixation) are proposed; high external N concentrations and foliar uptake of volatilised NH3. The leaf delta(15)N value of sugarcane grown in aerated solution culture was shown to decrease by approximately 5parts per thousand with increasing external N concentration (0.5-8.0 mM), with both NO3- and NH4+ nitrogen forms. Foliar uptake of atmospheric NH3 has been shown to result in depleted leaf delta(15)N values in many plant species. Acid traps collected atmospheric N with negative delta(15)N value (-24.45+/-0.90parts per thousand) from above a field recently surface fertilised with urea. The delta(15)N of leaves of sugarcane plants either growing directly in the soil or isolated from soil in pots dropped by 3.00parts per thousand in the same field after the fertiliser application. Both the high concentration of external N in the root zone (following the application of N-fertilisers) and/or subsequent foliar uptake of volatilised NH3 could have caused the depleted leaf delta(15)N values measured in the sugarcane crops at these sites.