Genetic variation for carbon isotope composition in honey mesquite (Prosopis glandulosa)

Carbon isotope composition (delta(13)C) is a useful surrogate for integrated, plant water-use efficiency (WUE) when measured on plants grown in a common environment. In a variety of species, genetic variation in delta(13)C has been linked to the distribution of genotypes across gradients in atmospheric and soil water. We examined genetic variation for delta(13)C in seedlings of honey mesquite (Prosopis glandulosa Torr.), an invasive grassland shrub that thrives in the southwestern USA. Fifteen maternal families, representing progeny of 15 adult trees, were studied in three common garden experiments in a greenhouse. The 990-km east-west transect along which the adult trees were located encompasses a wide precipitation gradient, and includes mesic grassland, semiarid grassland, and Chihuahuan desert ecosystems. Genetic variation for delta(13)C in mesquite was substantial, with the rank order of half-sib families based on delta(13)C relatively stable across experiments, which were conducted under different environmental conditions. Conversely, rankings of families by mean seedling height (an index of growth rate) varied markedly among experiments. Seedlings derived from Chihuahuan desert adults emerged more quickly and had more negative delta(13)C (indicative of lower WUE) than seedlings derived from the other regions. Although delta(13)C and seedling height were not correlated, these results suggest that mesquite genotypes at the drier, western extreme of the species' range are adapted for quicker emergence and possibly faster growth than genotypes from mesic areas. Together, these traits may facilitate exploitation of infrequent precipitation events.     

Physiological variation among Populus fremontii populations: short- and long-term relationships between delta13C and water availability

Different populations of widely distributed species can experience dramatically different climatic conditions that may influence physiological activity, specifically carbon assimilation and water use. Populus fremontii Wats. (Fremont cottonwood) populations are found near rivers of varying size along a precipitation gradient from New Mexico to northern California. Climatic differences among populations may lead to physiological differences because P. fremontii is sensitive to water availability. To assess physiological variation among populations, we collected foliage and wood samples from 13 populations that experience different precipitation and stream flow regimes and analyzed the samples for carbon isotope composition (delta13C). Wood delta13C served as a lifetime-averaged indicator of water-use efficiency (WUE), whereas foliage delta13C provided as an estimate of WUE during the growing season of collection. We found approximately 3.4 per thousand variation in delta13C among populations for both foliage (-31.1 to -27.9 per thousand) and wood (-28.3 to -24.7 per thousand). Wood delta13C was, on average, 2.8 per thousand more enriched than foliage. Some of the variation in wood delta13C can be explained by variation in elevation of the study sites. We constructed total precipitation and mean stream flow variables based on the length of the growing season at each study site and analyzed for a relationship between delta13C, precipitation and stream flow. A significant relationship between foliage delta13C and precipitation was found, but water availability did not explain a significant fraction of the variation in wood delta13C. The data suggest that water availability can account for some of the delta13C variation among populations but, given the large residual variances, other factors are important.     

Dry mass allocation, water use efficiency and delta(13)C in clones of Eucalyptus grandis, E. grandis x camaldulensis and E. grandis x nitens grown under two irrigation regimes

Clonal variation in water use efficiency (WUE), dry mass accumulation and allocation, and stable carbon isotope ratio (delta(13)C) of crude leaf fiber extracts was determined in six clones of Eucalyptus grandis W. Hill ex Maiden. grown for 16 months in field lysimeters in two soil water regimes. The relationships between delta(13)C and WUE calculated on the basis of leaf, harvestable stem, shoot and whole-plant dry mass accumulation were investigated. There was no clonal variation in dry mass accumulation but clonal allocation to roots, harvestable stems, branches and leaves varied. Water use efficiencies (mass of plant or plant part/water used over 16 months) differed significantly between clones. The clonal ranking of WUE varied depending on the units of dry mass accumulation used. Significant relationships between delta(13)C values and instantaneous water use efficiencies and ratios of internal leaf to ambient CO(2) concentrations were found only in the high soil water treatment. There were no relationships between delta(13)C values and whole-plant, shoot and harvestable stem water use efficiencies and soil water availability. Values of delta(13)C were negatively correlated with dry mass accumulation in the low soil water treatment. At the whole-plant level, WUE was positively correlated with dry mass accumulation in the high soil water treatment. We found significant differences in delta(13)C values between clones and the clonal rankings in delta(13)C and WUE were maintained in both soil water treatments.     

Comparison of water-use efficiency of seedlings from two sympatric oak species: genotype x environment interactions

Seedlings of two sympatric oak species, Quercus robur L. and Quercus petraea (Matt.) Liebl., were grown in common garden conditions to test for potential interspecific differences in intrinsic water-use efficiency (WUE). Intrinsic water-use efficiency was estimated based on carbon isotope composition of shoots (delta13C) and on gas exchange measurements (ratio of net CO2 assimilation rate to stomatal conductance (A/g(sw))). In addition, genotype x environment interactions were tested by subjecting the seedlings to four irradiance treatments (8, 18, 48 and 100% of incident solar irradiance) imposed by neutral shading nets, and, in the 100% irradiance treatment, two watering regimes. In all treatments, initial growth of Q. robur was faster than that of Q. petraea. In both species, there was a tight correlation between delta13C and A/g(sw). Intrinsic water-use efficiency increased with increasing irradiance (almost doubling from 8 to 100% irradiance), and this effect paralleled the increase in A with increasing irradiance. In full sun, WUE of Q. petraea seedlings was 10-15% higher than in Q. robur seedlings, with the difference attributable to a difference between the species in g(sw). The interspecific difference in WUE was maintained during drought, despite the appreciable increase in WUE and decrease in growth imposed by drought. No interspecific differences in WUE were observed at low irradiances, suggesting a strong genotype x environment interaction for WUE. These findings confirm the existence of interspecific genetic differences in WUE, but also show that there is large intraspecific variability and plasticity in WUE. The initially greater height and biomass increments in Q. robur seedlings illustrate the ability of this species to out-compete Q. petraea in the early stages of forest regeneration. For adult trees growing in closed canopies, the high WUE of Q. petraea may contribute significantly to its survival during dry years, whereas the low WUE of Q. robur may account for the frequently observed declines in adult trees of this species following drought.     

Summer precipitation influences the stable oxygen and carbon isotopic composition of tree-ring cellulose in Pinus ponderosa

The carbon and oxygen isotopic composition of tree-ring cellulose was examined in ponderosa pine (Pinus ponderosa Dougl.) trees in the western USA to study seasonal patterns of precipitation inputs. Two sites (California and Oregon) had minimal summer rainfall inputs, whereas a third site (Arizona) received as much as 70% of its annual precipitation during the summer months (North American monsoon). For the Arizona site, both the delta(18)O and delta(13)C values of latewood cellulose increased as the fraction of annual precipitation occurring in the summer (July through September) increased. There were no trends in latewood cellulose delta(18)O with the absolute amount of summer rain at any site. The delta(13)C composition of latewood cellulose declined with increasing total water year precipitation for all sites. Years with below-average total precipitation tended to have a higher proportion of their annual water inputs during the summer months. Relative humidity was negatively correlated with latewood cellulose delta(13)C at all sites. Trees at the Arizona site produced latewood cellulose that was significantly more enriched in (18)O compared with trees at the Oregon or California site, implying a greater reliance on an (18)O-enriched water source. Thus, tree-ring records of cellulose delta(18)O and delta(13)C may provide useful proxy information about seasonal precipitation inputs and the variability and intensity of the North American monsoon.     

Photosynthesis and water-use efficiency of sugar maple (Acer saccharum) in relation to pear thrips defoliation

An experimental introduction of pear thrips (Taeniothrips inconsequens Uzel), a major defoliator in sugar maple (Acer saccharum Marsh.) forests in northeastern North America, was conducted in a field plantation to determine if compensatory gas exchange occurs in response to feeding damage by this piercing-sucking insect. Sugar maple trees were enclosed in netting (167 micro m mesh) and pear thrips adults were introduced before leaf expansion in the spring. Pear thrips reduced whole-tree leaf area by approximately 23% and reduced leaf size (both mass and area) by 20% in the upper crown. Measurements of net CO(2) assimilation rate (A(net)) and stomatal conductance (g(s)) were made on tagged foliage that was later analyzed for stable carbon isotope composition (delta(13)C) to provide estimates of short- and long-term leaf water use efficiency (WUE). Pear thrips feeding reduced A(net) for fully expanded leaves by approximately 20%, although leaf chlorophyll content and leaf mass per unit area were apparently not affected. Comparison of A(net), g(s), instantaneous WUE and leaf delta(13)C between damaged and control trees as well as visibly undamaged versus moderately damaged foliage on pear thrips-infested trees indicated that there were no effects of pear thrips feeding damage on WUE or leaf delta(13)C. Long-term WUE among sugar maple trees in the field plantation, indicated by leaf delta(13)C analysis, was related to shorter-term estimates of leaf gas exchange behavior such as g(s) and calculated leaf intercellular CO(2) concentration (C(i)). We conclude that pear thrips feeding has no effect on leaf WUE, but at the defoliation levels in our experiment, it may reduce leaf A(net), as a result of direct tissue damage or through reduced g(s). Therefore, even small reductions in leaf A(net) by pear thrips feeding damage may have an important effect on the seasonal carbon balance of sugar maple when integrated over the entire growing season.     

Variation in branchlet delta13C in relation to branchlet nitrogen concentration and growth in 8-year-old hoop pine families (Araucaria cunninghamii) in subtropical Australia

Carbon isotope composition (delta13C) of branchlet tissue at nine canopy positions, and nitrogen concentration (N(mass)) at four canopy positions, were assessed in 8-year-old hoop pine (Araucaria cunninghamii Ait. ex D. Don) trees from 23 half-sib families, grown in six blocks of a progeny test in southeastern Queensland, Australia. There was considerable variation among sampling positions, families and blocks in both delta13C and N(mass). The delta13C was positively related to N(mass) only for samples from the upper outer crown (P < 0.005). Phenotypic correlations existed between tree growth and canopy delta13C. Branchlet delta13C of the inner and lower outer crown was positively related (P < 0.037) to tree height, but delta13C in branchlets of the upper outer crown was not related to tree height, or was related negatively (P < 0.045). There were significant differences in delta13C between hoop pine families for six canopy positions (upper canopy positions as well as lower canopy positions on the northern side), with heritabilities greater than 0.40. The significance of these findings is discussed in relation to water and light competition within the tree canopy of hoop pine.     

Effective rainfall seasons for interannual variation in δ13C and tree-ring width in early and late wood of Chinese pine and black locust on the Loess Plateau, China

We studied the importance of effective rainfall for interannual variation in water use efficiency (WUE) and tree-ring growth of Chinese pine (Pinus tabulaeformis Carr.) and black locust (Robinia pseudoacacia L.) by examining correlations of seasonal precipitation with annual values of stable carbon isotope ratio (δ¹³C) and tree-ring width in early and late wood. The correlations with precipitation were examined for each month and for periods of all possible lengths from 2 to 22 months starting from January of the previous year to October of the current year. The period with the highest correlation was adopted as the most effective rainfall season for interannual variations in WUE and tree-ring width. In early wood, precipitation during the dry season (October to May) before the growing season was negatively correlated with δ¹³C in pine trees and positively correlated with ring width in pine and locust trees. In late wood, rainfall during the growing season in the current year was negatively correlated with δ¹³C in pine and locust trees, and positively correlated with ring width in locust trees. Our results demonstrated the differences in the water use strategies of pine and locust trees. The δ¹³C in pines indicated higher WUE and more conservative water use than in locust trees. Precipitation during the dry season affected the interannual variation in WUE and tree-ring growth in pine and locust trees, indicating that rainfall during the dry season is important for carbon gain and tree-ring growth during the following growing season.     

Carbon isotope discrimination and oxygen isotope composition in clones of the F(1) hybrid between slash pine and Caribbean pine in relation to tree growth,water-use efficiency and foliar nutrient concentration

The objectives of this study were: (1) to examine how foliar carbon isotope discrimination (Delta) and oxygen isotope composition (delta(18)O) are related to tree growth, ash mineral nutrient concentration and foliar nutrient concentration in 7-year-old clones of the F(1) hybrid between slash pine (Pinus elliottii Engelm.) and Caribbean pine (P. caribaea var. hondurensis Barr. et Golf.) in subtropical Australia; and (2) to evaluate the potential of using foliar Delta, ash mineral nutrient concentration and delta(18)O measurements for selecting F(1) hybrid pine clones with high water-use efficiency (WUE) and growth potential. There were significant differences in tree growth, foliar Delta, delta(18)O and ash mineral nutrient concentration among the eight clones tested. Significant negative linear relationships existed between tree growth and Delta, extrapolating to zero growth at Delta = 24-30 per thousand. There were strong genetic correlations (r = -0.83 to -0.96) between Delta and tree growth, particularly tree height. Significant non-genetic correlations (r = -0.62 to -0.80) existed between Delta and foliar K concentration. Foliar delta(18)O, ash mineral nutrient concentration and foliar nutrient concentration were unrelated to tree growth. In the F(1) hybrid pine clones, variation in tree WUE, as reflected by Delta, was largely attributed to a genetic effect on leaf photosynthetic capacity rather than on stomatal conductance, as reflected by foliar delta(18)O.     

Age-related variations in delta(13)C of ecosystem respiration across a coniferous forest chronosequence in the Pacific Northwest

We tested the hypothesis that forest age influences the carbon isotope ratio (delta13C) of carbon reservoirs and CO2 at local and regional levels. Carbon isotope ratios of ecosystem respiration (delta13C(R)), soil respiration (delta13C(R-soil)), bulk needle tissue (delta13C(P)) and soil organic carbon (delta(13)C(SOC)) were measured in > 450-, 40- and 20-year-old temperate, mixed coniferous forests in southern Washington, USA. Values of delta13C(R), delta13C(R-soil), delta13C(P) and delta13C(SOC) showed consistent enrichment with increasing stand age. Between the youngest and oldest forests there was an approximately 1 per thousand enrichment in delta13C(P) (at similar canopy levels), delta13C(SOC) (throughout the soil column), delta13C(R-soil) (during the wet season) and delta13C(R) (during the dry season). Mean values of delta13C(R) were -25.9, -26.5 and -27.0 per thousand for the 450-, 40- and 20-year-old forests, respectively. Both delta13C(R-soil) and the difference between delta13C(R) and delta13C(R-soil) were more 13C enriched in older forests than in young forest: delta13C(R) - delta13C(R-soil) = 2.3, 1.1 and 0.5 per thousand for the 450-, 40- and 20-year-old forests, respectively. Values of delta(13)C(P) were proportionally more depleted relative to delta13C(R): delta13C(R) - delta13C(P) = 0.5, 2.2 and 2.5 per thousand for the 450-, 40- and 20-year-old forests, respectively. Values of delta13C(P) were most 13C-enriched at the top of the canopy and in the oldest forest regardless of season (overall values were -26.9, -28.7 and -29.4 per thousand for the 450-, 40- and 20-year-old forests, respectively). Values of delta13C(SOC) from shallow soil depths were similar to delta13C(P) values of upper- and mid-canopy needles. All delta13C data are consistent with the hypothesis that a decrease in stomatal conductance associated with decreased hydraulic conductance leads to increased CO2 diffusional limitations in older coniferous trees. The strong associations between delta13C(P) in needles with delta13C(R) and delta13C(R-soil) at the forest level suggest that 13C observations scale between leaf and ecosystem levels.     

Differences in delta13C and diameter growth among remnant Scots pine populations in Scotland

Published data suggest that differences in wood cellulose carbon isotope composition (delta13C) and xylem ring width among natural populations of Scots pine in Scotland (Pinus sylvestris L.) are attributable to the persistence of palaeotypes of various post-glacial migratory origins. We assessed differences in wood cellulose delta13C and ring width among Scottish Scots pine populations grown in a clone bank and in natural stands at various locations in northern and central Scotland. Ring width and wood cellulose delta13C varied significantly among natural stands. Potential water deficit was positively correlated with wood cellulose delta13C and xylem ring width in the natural stands. Neither wood cellulose delta13C nor xylem ring width of clone bank trees correlated with any climate variables at the sites from which the trees originated, indicating little adaptation to climate for these traits. Xylem ring width showed a site x population interaction for the growth sites (i.e., natural stands versus clone bank), but wood cellulose delta13C did not. These results suggest that climate variation in Scotland has not resulted in significant genetic variation in wood cellulose delta13C or xylem ring width in post-glacial populations.     

Carbon isotope variation in Douglas-fir foliage: improving the delta(13)C-climate relationship

The natural abundance of stable carbon isotopes in the annual rings of forest trees is used as a tracer of environmental changes such as climate and atmospheric pollution. Although tree-ring delta(13)C varies by about 2 per thousand from year to year, variability within the foliage can be as high as 6 per thousand. Recent studies have shown that branch length affects stomatal response, which influences the integrated foliar delta(13)C signal. To improve the ability of delta(13)C to predict climate differences, we examined the relationship between branch length and foliar delta(13)C in Pseudotsuga menziesii (Mirb.) Franco from four sites across a steep climate gradient in Oregon. The transect spanned the boundary between the ranges of the coastal variety, P. menziesii var. menziesii (three sites), and the Rocky Mountain variety, P. menziesii var. glauca (one site). At the most maritime site, branch length explained 76% of within-site variation of 5 per thousand, whereas at the harshest site, branch length accounted for only 15% of this variation. We considered the possibility that cavitation in the water-conducting xylem obscures the branch length effect in the harsher climates. Cavitation, as measured by dye perfusion, was most extensive at sites where the branch length effect in the coastal variety was weakest. Trees at the site with the most substantial cavitation displayed seasonal xylem refilling. Branch length standardization significantly improved the relationship between delta(13)C and climate. With standardization to constant length, delta(13)C values were significantly related to the degree that climatic variables, as modeled with a forest growth simulation model, constrain transpiration (R(2) = 0.69, P < 0.0001). Without standardization, the R(2) was 0.27. We conclude that sampling standard length branches or tree rings from trees of similar shape and size is desirable when seeking correlations between isotopic composition and climate.     

Impact of fertilization on tree-ring delta15N and delta13C in beech stands: a retrospective analysis

We studied the effects of two fertilization treatments (N and NPKCa) on wood nitrogen (N) isotope composition (delta(15)N), water-use efficiency (WUE) estimated by carbon isotope composition (delta(13)C) analyses, and ring width of trees in 80-year-old beech (Fagus sylvatica L.) stands in the forest of Fougères, western France. Four replicates were fertilized in two successive years (1973 and 1974), 20 years before core sampling. Unfertilized control trees displayed a decreasing delta(15)N trend with time. The N and NPKCa treatments both increased delta(15)N compared with the control treatment. Wood extraction by organic solvents enhanced the delta(15)N signal. Thus, N addition to the beech ecosystem, even in moderate amounts, could be traced back in tree-ring delta(15)N, suggesting that wood N isotope analysis is a promising tool for studying the long-term effects of N deposition on forests. Although WUE decreased for about 6 years after N fertilization, WUE in NPKCa-treated trees did not differ significantly from that in control trees. Results were similar whether based on cellulose or total wood delta(13)C analysis, suggesting that extraction of cellulose is not necessary when studying fertilization impacts on WUE. The NPKCa treatment had a large impact on radial growth, causing a significant long-lasting increase of 29% compared with the control treatment. Nitrogen alone did not change radial growth significantly.     

Leaf and wood carbon isotope ratios, specific leaf areas and wood growth of Eucalyptus species across a rainfall gradient in Australia

Leaves and samples of recent wood of Eucalyptus species were collected along a rainfall gradient parallel to the coast of Western Australia between Perth in the north and Walpole in the south and along a southwest to northeast transect from Walpole in southwestern Australia, to near Mount Olga in central Australia. The collection included 65 species of Eucalyptus sampled at 73 sites and many of the species were collected at several sites along the rainfall gradient. Specific leaf area (SLA) and isotopic ratio of 13C to 12C (delta 13C) of leaves that grew in 2002, and tree ring growth and delta 13C of individual cell layers of the wood were measured. Rainfall data were obtained from the Australian Bureau of Meteorology for 29 locations that represented one or a few closely located collection sites. Site-averaged data and species-specific values of delta 13C decreased with decreasing annual rainfall between 1200 and 300 mm at a rate of 1.63 per thousand per 1000 mm decrease in rainfall. Responses became variable in the low rainfall region (< 300 mm), with some species showing decreasing delta 13C with rainfall, whereas delta 13C increased or remained constant in other species. The range of delta 13C values in the low rainfall region was as large as the range observed at sites receiving > 300 mm of annual rainfall. Specific leaf area varied between 2 and 6 m2 kg(-1) and tended to increase with decreasing annual rainfall in some species, but not all, whereas delta 13C decreased with SLA. The relationship between delta 13C and SLA was highly species and soil-type specific. Leaf-area-based nitrogen (N) content varied between 2 and almost 6 g m(-2) and decreased with rainfall. Thus, thicker leaves were associated with higher N content and this compensated for the effect of drought on delta 13C. Nitrogen content was also related to soil type and species identity. Based on a linear mixed model, statistical analysis of the whole data set showed that 27% of the variation in delta 13C was associated with changes in SLA, 16% with soil type and only 1% with rainfall. Additionally, 21% was associated with species identity. For a subset of sites with > 300 mm rainfall, 43% of the variation was explained by SLA, 13% by soil type and only 3% by rainfall. The species effect decreased to 9% because there were fewer species in the subset of sites. The small effect of rainfall on delta 13C was further supported by a path analysis that yielded a standardized path coefficient of 0.38 for the effect of rainfall on SLA and -0.50 for the effect of SLA on delta 13C, but an insignificantly low standardized path coefficient of -0.05 for the direct effect of rainfall on delta 13C. Thus, in contrast to our hypothesis that delta 13C decreases with rainfall independent of soil type and species, we detected no statistically significant relationship between rainfall and delta 13C in leaves of trees growing at sites receiving < 300 mm of rainfall annually. Rainfall affected delta 13C indirectly through soil type (a surrogate for water-holding capacity) across the rainfall gradient. Annual tree rings are not clearly visible in evergreen Eucalyptus species, even in the seasonally cool climate of SW Australia. Generally, visible density transitions in the wood are related not to a strict annual cycle but to periods of growth associated mainly with rainfall. The relationship between delta 13C of leaves and the width of these stem increments was not statistically significant. Analysis of stem growth periods showed that delta 13C in wood responded to rainfall events, but carbohydrate storage and reallocation also affected the isotopic signature. Although delta 13C in wood of any one species varied over a range of 2 to 4 per thousand, there was a general relationship between delta 13C of the leaves and the annual range of delta 13C in wood. We conclude that species-specific traits are important in understanding the response of Eucalyptus to rainfall and that the diversity of the genus may reflect its response to the large climatic gradient in Australia and to the large annual and interannual variations in rainfall at any one location.     

Variation in seed and seedling traits and their relations to geo-climatic factors among populations in Yunnan Pine (<Emphasis Type="Italic">Pinus yunnanensis</Emphasis>)

Differences in seed germination and seedling growth among populations of Pinus yunnanensis Franch. were quantified to provide information for use in the regeneration, reforestation, and improvement of this species. Seed germination and seedling growth traits at the nursery stage were investigated at the population level. We evaluated relationships between pairs of measured traits as well as the association between measured traits and geo-climatic variables including longitude, latitude, elevation, temperature and precipitation. Seedling growth showed significant differences although a low magnitude of variation was found in the seed germination traits among the populations. Some populations with strong and stable growth (e.g., the Baoshan population) may be selected as superior seed sources for regeneration and reforestation. Germination percentage was negatively correlated with elevation, and positively with precipitation. In addition, germination index, vigor index and seedling height were positively correlated with precipitation. Seed germination index, cotyledon number and length, and seedling diameter were significantly and positively correlated with seed weight. Strong correlations between pairs of traits will be helpful in early evaluations for the selection of certain prominent traits.     

The relationship between the needle age and the growth rate in Scots pine (Pinus sylvestris): a retrospective analysis by needle trace method (NTM)

Variation in annual radial and height increments (RI and HI, respectively) as well as the number of needles belonging to different needle sets of successive annual shoots, determined by the needle trace method (NTM), was studied retrospectively in Scots pine stands, one in southern Finland and two in Estonia. The connection between different needle sets and tree growth was studied by correlation and covariance analyses. The numbers of needles of the three youngest needle sets were positively correlated with growth, whereas the number of needles older than 3 years had no significant correlation. The covariance analysis emphasized the negative relationship of older needle sets with tree growth. It is generally concluded that the two or three youngest needle sets are important and all older needles minimally contribute to growth, possibly because of their small quantity. Older needles may remain on the tree if assimilation is impeded for some reason, e.g. due to premature loss in the two youngest needle sets. The trees having attached old needles grew less than those that had already totally lost these needle sets.     

Water availability and branch length determine delta(13)C in foliage of Pinus pinaster

The stable carbon isotope composition (delta(13)C) of foliage integrates signals resulting from environmental and hydraulic constraints on water movement and photosynthesis. We used branch length as a simple predictor of hydraulic constraints to water fluxes and determined the response of delta(13)C to varying water availability. Foliage up to 6 years old was taken from Pinus pinaster Ait. trees growing at four sites differing in precipitation (P; 414-984 mm year(-1)) and potential evaporation (ET; 1091-1750 mm year(-1)). Branch length was the principal determinant of temporal trends in delta(13)C. The strong relationship between delta(13)C and branch length was a function of hydraulic conductance, which was negatively correlated with branch length (r(2) = 0.84). Variation in P and ET among sites was reflected in delta(13)C, which was negatively correlated with P/ET (r(2) = 0.66). However, this analysis was confounded by differences in branch length. If the effects of branch length on delta(13)C were first removed, then the 'residual' delta(13)C was more closely related to P/ET (r(2) = 0.99), highlighting the importance of accounting for variation in hydraulic constraints to water flux between sites and years. For plant species that exhibit considerable phenotypic plasticity in response to changes in environment (e.g., variation in leaf area, branch length and number, or stem form), the environmental effects on delta(13)C in foliage can only be reliably assessed if deconvoluted from hydraulic constraints.     

Age structure of wind-felled canopy trees for Siebold’s beech (<Emphasis Type="Italic">Fagus crenata</Emphasis>) in the northernmost population in Karibayama,Hokkaido

To determine the lifespan of Siebold’s beech (Fagus crenata), a dominant canopy tree species of the cool temperate forests of Japan, we estimated the age of 33 mature beech trees felled by a large typhoon in September 2004 at Karibayama, in southern Hokkaido. The study site lies within one of the northernmost distributions of Siebold’s beech populations in Japan. We counted tree rings from tree disks at a height of 1 m, and diameter growth was measured in annual increments. The sample age revealed two major peaks at 240 and 175 years. The diameter growth was good between 40 and 60 years of age and gradually decreased after 70 years of age. Annual diameter increments were noticeably small in good fruiting years, indicating a large allocation of annual carbon gains to the reproductive organs. Bud scale counts of juvenile trees revealed that the age at which the height reached 1 m was approximately 6–13 years. The lifespan of Siebold’s beech in the Karibayama population was therefore estimated to be approximately 250 years, an estimation that is higher than that previously reported for this species in the northernmost populations.     

Roles of soil chemistry and water availability in site-related delta(13)C 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.     

Branchlet nutrient concentration in hoop pine (Araucaria cunninghamii) relative to family,stable carbon and oxygen isotope ratios and growth rate in contrasting environments

Genetic variation in branchlet nutrient (N, P, K, Na, Ca, Mg, Mn and Fe) concentrations and mineral concentration (sum of branchlet P, K, Na, Ca, Mg, Mn and Fe concentrations) of 8-9-year-old hoop pine (Araucaria cunninghamii Ait. ex D. Don) half-sib families was assessed for four canopy positions at a wet site (23 families) and two canopy positions at an N- and water-limiting dry site (22 families) in relation to tree growth and associated branchlet carbon (delta13C) and oxygen (delta18O) isotope composition in southeast Queensland, Australia. Branchlet nutrient and mineral concentrations varied significantly among families and with canopy position and site. Depending on the canopy position sampled, the hoop pine family effect accounted for 0 to 13.8% of the total variation in branchlet N concentration, and for 0 to 30.3% of the total variation in branchlet mineral concentration at the wet site. The corresponding values for the family effect at the dry site were 0-13.3% for branchlet N concentration and 0-25.7% for branchlet mineral concentration. There were significant variations in branchlet P, K, Ca and Mg concentrations at both sites, and these variations differed with canopy position. Relationships between family means of branchlet N concentration and tree growth or delta13C or delta18O varied with canopy position at both sites. At the wet site, there were significant positive correlations between branchlet mineral concentration in the upper-outer or upper-inner canopy and tree height (r = 0.26 and 0.37, P < 0.01) and between branchlet mineral concentration and delta13C (r = 0.24, P < 0.01) in the upper-inner canopy, and a significant negative correlation between branchlet mineral concentration and delta13C (r = -0.21, P < 0.05) in the upper-outer canopy. At the dry site, branchlet mineral concentrations in the upper-inner and upper-outer canopy were significantly correlated with branchlet delta13C (r = -0.28 and -0.51, P < 0.01), and branchlet N concentration in the upper-inner canopy was significantly correlated with tree growth (r = 0.29, P < 0.01). A significant correlation between branchlet delta18O (an index of stomatal conductance) and branchlet mineral concentration at the dry site (r = 0.39, P = 0.020) indicated that stomatal conductance might be a factor regulating the variation in branchlet mineral concentration of the hoop pine families. Both branchlet N concentration and mineral concentration at particular canopy positions assist in selecting hoop pine families with improved tree growth and N- and water-use efficiency in environments where both N deficiency and a limited water supply are major factors affecting plantation productivity.