Ozone air pollution effects on tree-ring growth, delta(13)C, visible foliar injury and leaf gas exchange in three ozone-sensitive woody plant species

We assessed the effects of ambient tropospheric ozone on annual tree-ring growth, delta(13)C in the rings, leaf gas exchange and visible injury in three ozone-sensitive woody plant species in southern Switzerland. Seedlings of Populus nigra L., Viburnum lantana L. and Fraxinus excelsior L. were exposed to charcoal-filtered air (CF) and non-filtered air (NF) in open-top chambers, and to ambient air (AA) in open plots during the 2001 and 2002 growing seasons. Ambient ozone exposures in the region were sufficient to cause visible foliar injury, early leaf senescence and premature leaf loss in all species. Ozone had significant negative effects on net photosynthesis and stomatal conductance in all species in 2002 and in V. lantana and F. excelsior in 2001. Water-use efficiency decreased and intercellular CO(2) concentrations increased in all species in response to ozone in 2002 only. The width and delta(13)C of the 2001 and 2002 growth rings were measured for all species at the end of the 2002 growing season. Compared with CF seedlings, mean ring width in the AA and NF P. nigra seedlings was reduced by 52 and 46%, respectively, in 2002, whereas in V. lantana and F. excelsior, ring width showed no significant reductions in either year. Although delta(13)C was usually more negative in CF seedlings than in AA and NF seedlings, with the exception of F. excelsior in 2001, ozone effects on delta(13)C were significant only for V. lantana and P. nigra in 2001. Among species, P. nigra exhibited the greatest response to ozone for the measured parameters as well as the most severe foliar injury and was the only species to show a significant reduction in ring width in response to ozone exposure, despite significant negative ozone effects on leaf gas exchange and the development of visible foliar injury in V. lantana and F. excelsior. Thus, significant ozone-induced effects at the leaf level did not correspond to reduced tree-ring growth or increased delta(13)C in all species, indicating that the timing of ozone exposure and severity of leaf-level responses may be important in determining the sensitivity of tree productivity to ozone exposure.     

Variation in foliar delta(13)C values within the crowns of Pinus radiata trees

Although herbaceous species generally show little within plant variation in delta(13)C, trees show large spatial and temporal differences. We found that the aspect of exposure and branch length accounted for up to 6 per thousand delta(13)C difference within the foliage of individual trees of Pinus radiata D. Don. The foliage on branches 0.5 m in length was as much as 4 per thousand more depleted in (13)C than foliage on 10-m long branches, and an additional 2 per thousand more depleted on the shaded side than on the exposed side. We confirmed that on clear days, relative branch hydraulic conductivity, defined as the ratio of transpiration to the water potential gradient, was much higher in short branches than in long branches. Stomatal conductance remained high in foliage on short branches during the day, whereas it declined progressively in long-branch foliage under similar conditions. These differences were sufficient to explain the observed variation in delta(13)C in foliage on long and short branches.     

Influence of canopy position, needle age and season on the foliar gas exchange of Pinus canariensis

We investigated the seasonal variation in the gas exchange of current and 1-year-old needles in the upper sun and lower shade crown of adult Pinus canariensis trees. In general, current year needles displayed lower gas exchange rates than the 1-year-old needles. In both needle age classes, gas exchange was significantly lower in the shade than in the sun crown. However irrespective of crown position and needle age, maximum daily net photosynthesis, transpiration, and stomatal conductance for water vapour were generally higher during the wet and cold winter as compared to the dry and hot summer. These higher gas exchange values obtained during the cold and wet season can mainly be explained by higher soil-water availability and lower evaporative demand as compared to the warm and dry seaon. In addition, we also observed a displacement in the temperature optimum of net photosynthesis towards lower temperatures during the cold and wet season as compared to the warm and dry season. The observed gas exchange characteristics indicate a conservative water saving strategy and thus allowing P. canariensis needles to maintain a positive carbon gain even at periods of high evaporative demand and low soil-water availability.     

Responses of foliar gas exchange to long-term elevated CO(2) concentrations in mature loblolly pine trees

Branches of field-grown mature loblolly pine (Pinus taeda L.) trees were exposed for 2 years (1992 and 1993) to ambient or elevated CO(2) concentrations (ambient + 165 micro mol mol(-1) or ambient + 330 micro mol mol(-1) CO(2)). Exposure to elevated CO(2) concentrations enhanced rates of net photosynthesis (P(n)) by 53-111% compared to P(n) of foliage exposed to ambient CO(2). At the same CO(2) measurement concentration, the ratio of intercellular to atmospheric CO(2) concentration (C(i)/C(a)) and stomatal conductance to water vapor did not differ among foliage grown in an ambient or enriched CO(2) concentration. Analysis of the relationship between P(n) and C(i) indicated no significant change in carboxylation efficiency of ribulose-1,5-bisphosphate carboxylase/oxygenase during growth in elevated CO(2) concentrations. Based on estimates derived from P(n)/C(i) curves, there were no apparent treatment differences in dark respiration, CO(2) compensation point or P(n) at the mean C(i). In 1992, foliage in the three CO(2) treatments yielded similar estimates of CO(2)-saturated P(n) (P(max)), whereas in 1993, estimates of P(max) were higher for branches grown in elevated CO(2) than in ambient CO(2). We conclude that field-grown loblolly pine trees do not exhibit downward acclimation of leaf-level photosynthesis in their long-term response to elevated CO(2) concentrations.     

Variability in growth, carbon isotope composition, leaf gas exchange and hydraulic traits in the eastern Mediterranean cedars Cedrus libani and C. brevifolia

Four Turkish provenances and five Lebanese provenances of Cedrus libani A. Rich. and one Cypriot provenance of C. brevifolia Henry were compared during the third year of growth in a controlled-climate greenhouse after exposure to a well-watered or moderate-drought treatment. Effects of treatment on CO(2) assimilation (A), stomatal conductance (g(s)), (13)C isotope composition (delta(13)C), growth and biomass were assessed. Hydraulic conductivity and shoot vulnerability to cavitation were measured in well-watered plants only. The Lebanese provenances of C. libani had the highest growth rates, but were the most sensitive to drought. The Turkish provenances of C. libani showed moderate growth rates and moderate drought sensitivity. Cedrus brevifolia had the lowest growth rate and was least sensitive to drought. For each provenance, mean biomass values were positively correlated with delta(13)C and intrinsic water-use efficiency (A/g(s)), and negatively correlated with g(s). Drought reduced growth and favored carbon storage in roots, increasing the ratio of root biomass to aboveground biomass. The drought treatment increased delta(13)C and A/g(s). Specific hydraulic conductivity (K(s)) was similar for the provenance groups, whereas leaf-specific conductivity (K(l)) was lower in the Lebanese provenances than in the other provenances. Within each provenance group, provenances with the highest K(l) were most susceptible to xylem cavitation, but were also the most productive. Growth and drought adaptation were linked with precipitation in each provenance's native range.     

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.     

Effects of branch height on leaf gas exchange,branch hydraulic conductance and branch sap flux in open-grown ponderosa pine

Recent studies have shown that stomata respond to changes in hydraulic conductance of the flow path from soil to leaf. In open-grown tall trees, branches of different heights may have different hydraulic conductances because of differences in path length and growth. We determined if leaf gas exchange, branch sap flux, leaf specific hydraulic conductance, foliar carbon isotope composition (delta13C) and ratios of leaf area to sapwood area within branches were dependent on branch height (10 and 25 m) within the crowns of four open-grown ponderosa pine (Pinus ponderosa Laws.) trees. We found no difference in leaf gas exchange or leaf specific hydraulic conductance from soil to leaf between the upper and lower canopy of our study trees. Branch sap flux per unit leaf area and per unit sapwood area did not differ between the 10- and 25-m canopy positions; however, branch sap flux per unit sapwood area at the 25-m position had consistently lower values. Branches at the 25-m canopy position had lower leaf to sapwood area ratios (0.17 m2 cm-2) compared with branches at the 10-m position (0.27 m2 cm-2) (P = 0.03). Leaf specific conductance of branches in the upper crown did not differ from that in the lower crown. Other studies at our site indicate lower hydraulic conductance, sap flux, whole-tree canopy conductance and photosynthesis in old trees compared with young trees. This study suggests that height alone may not explain these differences.     

Responses of decomposition rate, nutrient return, and composition of leaf litter to thinning intensities in a Pinus tabulaeformis plantation

It is important to study the effect of tree density on the substrate quality and decomposition rate of leaf litter in plantations. In 2002, an experiment of the effects of thinning intensities at four different levels (i.e., 0 (I), 35.7% (II), 49.2% (III), and 64.2% (IV)) on undergrowth were carried out in an 18-year-old Pinus tabulaeformis plantation at an initial density of 3130 trees/hm² in the middle of the hills of Yingpan, Yanqing County, Beijing. Three years later, the rates of decomposition, the amount of nutrients returned, and the characteristics of leaf litter were compared by a litter bag method. The results show that the annual loss of dry matter of leaf litter in plots I, II, III, and IV was 25.81%, 26.25%, 27.68%, and 25.96%, respectively. The turnover of leaf litter was 10.04, 9.84, 9.24, and 9.97 years, respectively. Therefore, it is feasible and convenient to evaluate the effect of thinning on the rate of decomposition. In the first two months, the N, P, K, and Mg nutrients were released quickly. During the entire observation period of 14 months, the return of both N and Mg nutrients in the four plots exhibited a release-accumulation type of cycle. The return of P and K indicated a trend of release-accumulation and a relative balance. However, the return of Ca was far different from that of the other nutrients in the four plots. The total nutrient return of the four elements N, P, K, and Mg in the plots was 10.806, 31.016, 31.798, and 39.365 g/kg, respectively. Specifically, the quality of leaf litter in plot I was the worst in that N and Ca accumulated only 2.567 and 0.767 g/kg, respectively. Thinning did decrease the ratios of lignin to N and C to N and accelerated the rate of decomposition of leaf litter. The content of crude ash in leaf litter was enhanced by thinning, which prevented acid material, such as tannins and resins, from returning to the soil. The effect of thinning intensity was evaluated by the ratio of lignin to N, the ratio of C to N, and the ash content in leaf litter.     

Growth,morphology and gas exchange responses of two-year-old Quercus castaneifolia seedlings to flooding stress

Tolerance to flooding is crucial when thinking in promissory species for restoration of ecosystems prone to suffer soil water excess. In this study, we tested the flooding tolerance of two-years-old seedlings of Chestnut-leaved oak (Quercus castaneifolia C.A.Mey.) to determine whether it can be recommended for use in wetland restoration programs. Seedlings of Q. castaneifolia were subjected to three treatments: (1) control (C), (2) flooding for 60 days followed by a 42-day recovery period (F?+?R) and (3) continuous flooding for 102 days (F). Physiological performance, plant morphological changes and biomass accumulation were assessed. Results showed that, although net photosynthetic rates, stomatal conductance and transpiration decreased with prolonged flooding, when flood waters were removed, plants were able to recover their physiological activity (49–80% compared to controls). By contrast, when plants were continually flooded, their physiological activity decreased as well as the leaves experienced precocious senescence and wilting. Biomass responses paralleled physiological responses: leaf and root biomass were 42–49% higher under F?+?R treatment than under continuous flooding, and all plants under the F?+?R survived. Therefore, Q. castaneifolia appears as a promising species to be further studied when thinking re-vegetation of riverine areas and other temporarily flooded wetlands.     

Branch growth and gas exchange in 13-year-old loblolly pine (Pinus taeda) trees in response to elevated carbon dioxide concentration and fertilization

We used whole-tree, open-top chambers to expose 13-year-old loblolly pine (Pinus taeda L.) trees, growing in soil with high or low nutrient availability, to either ambient or elevated (ambient + 200 micromol mol-1) carbon dioxide concentration ([CO2]) for 28 months. Branch growth and morphology, foliar chemistry and gas exchange characteristics were measured periodically in the upper, middle and lower crown during the 2 years of exposure. Fertilization and elevated [CO2] increased branch leaf area by 38 and 13%, respectively, and the combined effects were additive. Fertilization and elevated [CO2] differentially altered needle lengths, number of fascicles and flush length such that flush density (leaf area/flush length) increased with improved nutrition but decreased in response to elevated [CO2]. These results suggest that changes in nitrogen availability and atmospheric [CO2] may alter canopy structure, resulting in greater foliage retention and deeper crowns in loblolly pine forests. Fertilization increased foliar nitrogen concentration (N(M)), but had no consistent effect on foliar leaf mass (W(A)) or light-saturated net photosynthesis (A(sat)). However, the correlation between A(sat) and leaf nitrogen per unit area (N(A) = W(A)N(M)) ranged from strong to weak depending on the time of year, possibly reflecting seasonal shifts in the form and pools of leaf nitrogen. Elevated [CO2] had no effect on W(A), N(M) or N(A), but increased A(sat) on average by 82%. Elevated [CO2] also increased photosynthetic quantum efficiency and lowered the light compensation point, but had no effect on the photosynthetic response to intercellular [CO2], hence there was no acclimation to elevated [CO2]. Daily photosynthetic photon flux density at the upper, middle and lower canopy position was 60, 54 and 33%, respectively, of full sun incident to the top of the canopy. Despite the relatively high light penetration, W(A), N(A), A(sat) and R(d) decreased with crown depth. Although growth enhancement in response to elevated [CO2] was dependent on fertilization, [CO2] by fertilization interactions and treatment by canopy position interactions generally had little effect on the physiological parameters measured.     

Variations in growth, survival and carbon isotope composition (delta(13)C) among Pinus pinaster populations of different geographic origins

To evaluate differences in growth and adaptability of maritime pine (Pinus pinaster Ait.), we studied growth, polycyclism, needle tissue carbon isotope composition (delta(13)C) as an estimate of water-use efficiency (WUE) and survival of seven populations at 10 years of age growing in a performance trial at a provenance test site in Escaroupim, Portugal. Six populations were from relatively high rainfall sites in Portugal and southwestern France (Atlantic group), and one population was from a more arid Mediterranean site in Spain. There were significant differences between some populations in total height, diameter at breast height, delta(13)C of bulk needle tissue, polycyclism and survival. A population from central Portugal (Leiria, on the Atlantic coast) was the tallest and had the lowest delta(13)C. Overall, the variation in delta(13)C was better explained by the mean minimum temperatures of the coldest month than by annual precipitation at the place of origin. Analyses of the relationships between delta(13)C and growth or survival revealed a distinct pattern for the Mediterranean population, with low delta(13)C (and WUE) associated with the lowest growth potential and reduced survival. There were significant negative correlations between delta(13)C and height or survival in the Atlantic group. Variation in polycyclism was correlated with annual precipitation at the place of origin. Some Atlantic populations maintained a high growth potential while experiencing moderate water stress. A detailed knowledge of the relationships between growth, survival and delta(13)C in contrasting environments will enhance our ability to select populations for forestry or conservation.     

Family variation in gas exchange, growth and leaf traits of black locust half-sib families

Variations in several growth, gas exchange and leaf traits among greenhouse-grown black locust (Robinia pseudoacacia L.) seedlings from 11 half-sib families were investigated. Three weeks after germination, early growth rates ranged from a minimum of 1 cm d(-1) in the slowest growing family, to a maximum of 3 cm d(-1) in the fastest growing family. Significant family variation in net photosynthetic rate per unit leaf area (P(N)), stomatal conductance, chlorophyll content, height, diameter, stem dry weight and total dry weight was observed. Net photosynthetic rate declined with seedling age. Net photosynthetic rate per unit leaf area was significantly correlated (r < 0.4) with specific leaf area, total chlorophyll, root dry weight, foliage dry weight and total dry weight. The correlation coefficients were higher (r >/= 0.55) between P(N) x total leaf area and growth traits (height, stem dry weight, foliage dry weight and total dry weight). The study indicated that variation in leaf area among the families was one reason for the lack of a strong relationship between P(N) and growth.     

Leaf water potential and gas exchange of eucalypt clonal seedlings to leaf solar protectant

This experiment was carried out in acclimatized greenhouses with seedlings of two hybrid clones of Eucalyptus urophylla×Eucalyptus grandis. A sunscreen protector consisting of 62.5% calcium carbonate was sprayed on the seedlings at weekly intervals. Water stress was induced by suspending irrigation until the soil reached 30% available water and water was then replaced so that it returned to field capacity. Gas exchange and leaf water status were measured after 50 days. The experiment was set up in a 4×2 factorial randomized block design in four distinct environments:(1) temperatures less than 21.2℃ and vapor pressure deficit of 0.15 kPa;(2) intermediate temperatures of 24.2℃ and vapor pressure deficit of 0.69 kPa;(3) high temperatures of 27.0℃ and high vapor pressure deficit of 1.4 kPa; and,(4) high temperature of 27.0℃ and vapor pressure deficit below 1.10 kPa. Two leaf sun protector treatments were used, with five replications each. High atmospheric demand acted as a stress factor for the seedlings during the initial growth phase.Applications of leaf sunscreen protector provided beneficial effects in maintaining optimum water status and gas exchanges of the plants under water stress.     

Responses of gas exchange to reversible changes in whole-plant transpiration rate in two conifer species

This study examined the autonomy of branches with respect to the control of transpiration (E) in Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and western red cedar (Thuja plicata Donn) seedlings. Experiments were conducted on whole seedlings in a gas exchange system with a dual-cuvette that permitted independent manipulation and measurement of E in the upper and lower cuvettes. The value of E in one cuvette was manipulated by varying vapor pressure deficit (D) between 2.2 and 0.2 kPa, whereas D in the other cuvette was held at 2.2 kPa. Reducing D, while increasing stomatal conductance (gs), resulted in an overall decrease in E. In western red cedar, this decrease was almost threefold, and in Douglas-fir, approximately fourfold. In well-watered western red cedar, a reduction of whole-plant E by 46% (brought about by reducing D in the upper cuvette) resulted in a 12% increase in gs, a 12% increase in E and a 7% increase in net assimilation (A) of untreated foliage in the lower cuvette. Responses of gs, E and A of untreated foliage were similar irrespective of whether foliage was at the top or bottom of the seedling. When D in the treatment cuvette was restored to 2.2 kPa, gs, E and A of foliage in the untreated cuvette returned to pretreatment values. In contrast, in well-watered Douglas-fir, there was almost no change in gs, E or A of untreated foliage in one cuvette when D in the other cuvette was reduced, causing a 52% reduction in whole-plant E. However, similar manipulations on drought-stressed Douglas-fir led to 7-19% increases in gs, E and A of untreated foliage. In well-watered western red cedar, daytime leaf water potential (Psil) was maintained near -0.9 MPa over a wide range of D, whereas Psil of Douglas-fir decreased from -1.2 to -1.5 MPa as D increased. The tighter (isohydric) regulation of Psil in western red cedar may partly explain its greater stomatal response to D and variation in whole-plant E compared with Douglas-fir. In response to a reduction in E, measured increases in Psil and gs of unmanipulated foliage were less than predicted by a model assuming complete hydraulic connectivity of foliage. Our results suggest the foliage of both species is partially autonomous with respect to water.     

Responses of Pinus massoniana and Pinus taeda to freezing in temperate forests in central China

Abstract

Masson pine (Pinus massoniana Lamb.), a native species widely distributed in temperate forests in central China, and Loblolly pine (Pinus taeda L.), an exotic tree species introduced to China from southeastern United States, are dominant evergreen conifers that play a pivotal role in maintaining forest structure and functions for the region. We examined the effects of freezing on these species with chlorophyll fluorescence and electrolyte leakage using both field- and laboratory-based experiments in September 2009 and January 2010, respectively. We found that freezing could cause a greater impact on the Loblolly pine than the Masson pine. Although the two species showed similar values of F _v /F _m and electrolyte leakage before freezing, the Masson pine needles showed lower F _v /F _m and higher electrolyte leakage ratios than those of the Loblolly pine when treated in low temperatures (?15 to 0°C). We also found that cold-acclimation was crucial for both species to adapt to low temperatures with the F _v /F _m ratio decreased approximately by 80% in the first freezing hour for the non-acclimated needles of both species while the cold-acclimated needles showed little changes in the F _v /F _m ratio. This finding is also supported by our measurements of electrolyte leakage. These results suggest that the Loblolly pine could be more susceptible to freezing damages than the Masson pine in central China.     

Seasonal patterns of leaf gas exchange and water relations in dry rain forest trees of contrasting leaf phenology

Diurnal and seasonal patterns of leaf gas exchange and water relations were examined in tree species of contrasting leaf phenology growing in a seasonally dry tropical rain forest in north-eastern Australia. Two drought-deciduous species, Brachychiton australis (Schott and Endl.) A. Terracc. and Cochlospermum gillivraei Benth., and two evergreen species, Alphitonia excelsa (Fenzal) Benth. and Austromyrtus bidwillii (Benth.) Burret. were studied. The deciduous species had higher specific leaf areas and maximum photosynthetic rates per leaf dry mass in the wet season than the evergreens. During the transition from wet season to dry season, total canopy area was reduced by 70-90% in the deciduous species and stomatal conductance (g(s)) and assimilation rate (A) were markedly lower in the remaining leaves. Deciduous species maintained daytime leaf water potentials (Psi(L)) at close to or above wet season values by a combination of stomatal regulation and reduction in leaf area. Thus, the timing of leaf drop in deciduous species was not associated with large negative values of daytime Psi(L) (greater than -1.6 MPa) or predawn Psi(L) (greater than -1.0 MPa). The deciduous species appeared sensitive to small perturbations in soil and leaf water status that signalled the onset of drought. The evergreen species were less sensitive to the onset of drought and g(s) values were not significantly lower during the transitional period. In the dry season, the evergreen species maintained their canopies despite increasing water-stress; however, unlike Eucalyptus species from northern Australian savannas, A and g(s) were significantly lower than wet season values.     

Responses of Picea, Pinus and Pseudotsuga roots to heterogeneous nutrient distribution in soil

The spatial distribution of plant-available mineral nutrients in forest soils is often highly heterogeneous. To test the hypothesis that local nutrient enrichment of soil leads to increased root proliferation in the nutrient-rich soil zone, we studied the effects of nutrient enrichment on the growth and nutrient concentrations of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) roots. Three-year-old seedlings were grown for 9 months in split-root containers filled with nutrient-poor forest mineral soil, with one side supplemented with additional mineral nutrients. Root dry weight and root length in Scots pine and Norway spruce were increased in the nutrient-supplemented soil compared with the nonsupplemented side, whereas root growth in Douglas-fir was unaffected by nutrient enrichment. Of the three species examined, Norway spruce exhibited the highest root and shoot growth and the highest nutrient demand. Specific root length (m g(-1)) and the number of root tips per unit root length were not affected by local nutrient addition in any of the species. Despite increased root growth in Norway spruce and Scots pine in nutrient-supplemented soil, their root systems contained similar nutrient concentrations on both sides of the split-root container. Thus, coniferous trees may respond to local nutrient supply by increased root proliferation, but the response varies depending on the species, and may only occur when trees are nutrient deficient. As a response to local nutrient enrichment, increases in root dry matter or root length may be better indicators of pre-existing nutrient deficiencies in conifers than increases in root nutrient concentrations.     

Effects of age-related increases in sapwood area,leaf area,and xylem conductivity on height-related hydraulic costs in two contrasting coniferous species

Introduction  

Knowledge of vertical variation in hydraulic parameters would improve our understanding of individual trunk functioning and likely have important implications for modeling water movement to the leaves. Specifically, understanding how foliage area (A _l), sapwood area (A _s), and hydraulic specific conductivity (k _s) vary with canopy position to affect leaf-specific conductivity (LSC) and whole-tree leaf-specific hydraulic conductance (K _l) may explain some of the contrasting patterns of A _l/A _s reported in the literature.     

A multi-species comparison of delta13C from whole wood, extractive-free wood and holocellulose

The stable carbon (C) isotope composition (delta13C) of tree rings is a powerful metric for reconstructing past physiological responses to climate variation. However, accurate measurement and interpretation are complicated by diagenesis and the translocation of compounds with distinct isotopic signatures. Isolation and analysis of cellulose minimizes these complications by eliminating variation due to biosynthetic pathways; however, isolation of cellulose is time-consuming and has no clear endpoint. A faster and better-defined analytical method is desirable. Our objectives were to determine if there is a direct relationship between the isotopic compositions of whole wood (WW), whole wood treated with solvents to remove mobile extractives (extractive-free wood; EF) and holocellulose (HC) isolated by extractive removal and subsequent bleaching. We also determined if total C concentration could explain the isotopic composition and variation among these three wood components of each sample. A set of wood samples of diverse phylogeny, anatomy and chemical composition, was examined. The mean offset or difference between HC and EF delta13C was 1.07 +/- 0.09 per thousand and the offset between HC and WW was 1.32 +/- 0.10 per thousand. Equivalence tests (with alpha = 0.05) indicated that the relationship between EF delta13C and HC delta13C had a slope significantly similar to 1 +/- 5.5%, whereas for the WW delta13C: HC delta13C relationship, the slope was significantly similar to 1 +/- 10.08%. A regression model using EF delta13C to predict HC delta13C had a slope of 0.97, which was not significantly different from unity (P = 0.264), whereas the regression for WW had a slope of 0.92 which was significantly different from unity (P = 0.0098). Carbon concentration was correlated with HC:WW offset and cellulose:EF offset (P = 0.0501 and 0.007, respectively), but neither relationship explained much of the variation (r2 = 0.12 and 0.14, respectively). We suggest that HC extraction is unnecessary for most analyses of tree-ring delta13C; a simple solvent extraction is a suitable alternative for many applications.     

A comparison of three approaches to modeling leaf gas exchange in annually drought-stressed ponderosa pine forests

We tested, compared and modified three models of stomatal conductance at the leaf level in a forest ecosystem where drought stress is a major factor controlling forest productivity. The models were tested against 2 years (1999 and 2000) of leaf-level measurements on ponderosa pine (Pinus ponderosa Dougl. ex Laws.) growing in the Mediterranean climate of California, USA. The Ball, Woodrow and Berry (1987) (BWB) model was modified to account for soil water stress. Among the models, results of the modified BWB model were in the closest agreement with observations (r2 = 0.71). The Jarvis (1976) model showed systematic simulation errors related to vapor pressure deficit (r2 = 0.65). Results of the Williams, Rastetter, Fernandes et al. (1996) (SPA) model showed the poorest correlation with empirical data, but this model has only one calibration parameter (r2 = 0.60). Sensitivity analyses showed that, in all three models, predictions of stomatal conductance were most responsive to photosynthetically active radiation and soil water content. Stomatal conductance showed little sensitivity to vapor pressure deficit in the Jarvis model, whereas in both the BWB and SPA models, vapor pressure deficit (or relative humidity) was the third most important variable. Parameterization of the SPA model was in accordance with the parameterization of the modified BWB model, although the two models differ greatly. Measured and modeled results indicate that stomatal behavior is not water conservative during spring; however, during summer, when soil water content is low and vapor pressure deficit is high, stomatal conductance decreases and, according to the models, intrinsic water- use efficiency increases.