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PubMed2001

Photosynthesis and light-use efficiency by plants in a Canadian boreal forest ecosystem.

Measurements of the photosynthetic response to midsummer irradiance were made for 11 species representing the dominant trees, understory shrubs, herbaceous plants and moss species in an old black spruce (Picea mariana (Mill.) B.S.P.) boreal forest ecosystem. Maximum rates of photosynthesis per unit foliage area at saturating irradiance, A(max), were highest for aspen (Populus tremuloides Michx.), reaching 16 micromol m(-2) s(-1). For tamarack (Larix laricina (Du Roi) K. Kock) and P. mariana, Amax was only 2.6 and 1.8 micromol m(-2) s(-1), respectively. Values of A(max) for understory shrubs and herbaceous plants were clustered between 9 and 11 micromol m(-2) s(-1), whereas A(max) of feather moss (Pleurozium schreberi (Brid.) Mitt.) reached only 1.9 micromol m(-2) s(-1). No corrections were made for differences in shoot structure, but values of photosynthetic light-use efficiency were similar for most species (70-80 mmol CO2 mol(-1)); however, they were much lower for L. laricina and P. mariana (15 mmol CO2 mol(-1)) and much higher for P. schreberi (102 m;mol CO2 mol(-1)). There was a linear relationship between Amax and foliage nitrogen concentration on an area basis for the broad-leaved species in the canopy and understory, but the data for P. mariana, L. laricina and P. schreberi fell well below this line. We conclude that it is not possible to scale photosynthesis from leaves to the canopy in this ecosystem based on a single relationship between photosynthetic rate and foliage nitrogen concentration.

Ecosystem↗
PubMed2002

Measured and predicted changes in tree and stand water use following high-intensity thinning of an 8-year-old Eucalyptus nitens plantation.

We investigated changes in the pattern of water use of an 8-year-old Eucalyptus nitens (Deane and Maiden) Maiden plantation soon after thinning. Sap flow sensors using heat pulse technology were deployed across three stands thinned to a final density of 100, 250 or 600 trees ha-1 plus an unthinned control (1250 trees ha-1). Changes in the relationship between tree size and daily water use were measured for 4 to 7 months after thinning. Thinning had no effect on sapwood water content. The increase in tree water use as a result of thinning was driven largely by significant changes in the radial pattern of sap velocity through the sapwood. The use of a canopy fraction factor in the Penman-Monteith equation to account for discontinuous canopies showed promise as a simple and effective method of scaling the model to predict transpiration from thinned plantations.

Circadian Rhythm↗
PubMed2002

Use of a physiological process model with forestry yield tables to set limits on annual carbon balances.

We present an approach that sets limits on annual carbon fluxes for different aged forests by using a simple process-based model (3-PG) and information derived from yield tables and local weather stations. Given a measure of height-growth potential, model predictions are constrained to match stand dynamics described in yield tables. Thus constrained, the model can provide reasonable annual estimates of gross photosynthesis under a specified climate, even with inexact knowledge of soil properties. If we assume that leaf litterfall and fine-root turnover approach equilibrium at canopy closure, maximum net annual ecosystem exchange can also be predicted from modeled estimates of these two detrital components and estimates of foliage, branch, stem and coarse-root production. The latter four components of production are predicted from allometric relationships with mean stem diameter. The approach is demonstrated for Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) stands between Ages 20 and 150 years growing under conditions typical of those at Wind River, Washington, USA. Gross photosynthesis (Pg) by Douglas-fir at Ages 20, 70 and 150 years with leaf area indices (L) of 8.1, 6.9 and 4.0 was predicted at 1630, 1580 and 1160 g C m-2 year(1, respectively. Maximum net ecosystem production (Pe) for the same range in age classes was predicted to average 275, 294 and 207 g C m-2 year-1, respectively. The predicted reductions in L for older stands do not occur because other species fill the canopy gaps created by natural mortality of Douglas-fir. As a result of the development of an understory, total Pg is predicted to decrease only slightly with the aging of the overstory. Estimates of Pe exclude respiration from coarse woody debris, although additions of this component are provided annually by the model. The process-based modeling approach, constrained by yield table estimates of stand properties, sets reasonable limits on annual carbon exchange and suggests which environmental variables deserve careful monitoring to refine estimates of carbon fluxes.

Carbon↗
PubMed2002

Water limitations to carbon exchange in old-growth and young ponderosa pine stands.

We investigated the impact of seasonal soil water deficit on the processes driving net ecosystem exchange of carbon (NEE) in old-growth and recently regenerating ponderosa pine (Pinus ponderosa Doug. ex Laws.) stands in Oregon. We measured seasonal patterns of transpiration, canopy conductance and NEE, as well as soil water, soil temperature and soil respiration. The old-growth stand (O) included two primary age classes (50 and 250 years), had a leaf area index (LAI) of 2.1 and had never been logged. The recently regenerating stand (Y) consisted predominantly of 14-year-old ponderosa pine with an LAI of 1.0. Both stands experienced similar meteorological conditions with moderately cold wet winters and hot dry summers. By August, soil volumetric water content within the upper 30 cm had declined to a seasonal minimum of 0.07 at both sites. Between April and June, both stands showed similar rates of transpiration peaking at 0.96 mm day(-1); thereafter, trees at the Y site showed increasing drought stress with canopy stomatal resistance increasing 6-fold by mid-August relative to values for trees at the O site. Over the same period, predawn water potential (psi(pd)) of trees at the Y site declined from -0.54 to -1.24 MPa, whereas psi(pd) of trees at the O site remained greater than -0.8 MPa throughout the season. Soil respiration at the O site showed a strong seasonal correlation with soil temperature with no discernible constraints imposed by declining soil water. In contrast, soil respiration at the Y site peaked before seasonal maximal soil temperatures and declined thereafter with declining soil water. No pronounced seasonal pattern in daytime NEE was observed at either site between April and September. At the Y site this behavior was driven by concurrent soil water limitations on soil respiration and assimilation, whereas there was no evidence of seasonal soil water limitations on either process at the O site.

Carbon↗
PubMed2003

Predicting leaf area index from scaling principles: corroboration and consequences.

Leaf area index (LAI) is a key biophysical variable in most process-based forest-ecosystem models. However, most such models require LAI as an input, typically obtained from empirical observations. We tested whether scaling principles based on trade-offs between single leaf and canopy properties could be effectively used to model LAI, thereby obviating the need for empirical observations. To do so, we used the process-oriented model, PnET, configured to estimate LAI from these same scaling principles. We derived biologically based LAI predictions (LAIPnET) for the Harvard Forest (Massachusetts, USA) eddy covariance tower site, a predominately mixed deciduous hardwood forest, using PnET, and compared these with a locally observed phenology record and with LAI estimates from both local (ground-based) photosynthetically active radiation transmittance (LAITRANS) and normalized difference vegetation index satellite data (LAINDVI). We generated the LAIPnET trajectory by running the PnET model with meteorological observations from the flux tower as model drivers. We derived LAITRANS from measurements of above- and below-canopy photosynthetically active radiation at the flux tower, and LAINDVI from observations from the Advanced Very High Resolution Radiometer (AVHRR) satellite-borne sensor of surface greenness for the 1 km2 cell containing the flux tower. Over a 5-year period, LAIPnET and LAITRANS values were comparable intra- and interannually, with maximum values differing by less than 0.1 to 0.2 LAI units (m2 m(-2)). Values of LAINDVI were similar to LAIPnET and LAITRANS in midsummer, but higher LAI values were predicted in the early and late portions of the growing season. In addition, we used the three alternative LAI trajectories in a modified version of the PnET model and compared the resulting outputs of gross primary production (GPP) with GPP estimates from the flux tower for 5 continuous years. The LAIPnET and LAITRANS inputs resulted in a difference of less than 3% in mean annual GPP from 1995 to 1999, and these were within 7 and 9%, respectively, of the annual eddy flux-based estimates over the same time period. The results indicate that biologically based LAI scaling approaches can closely track temporal changes in a deciduous forest and have potential for spatial and temporal scaling of LAI.

Ecosystem↗
PubMed2004

Component carbon fluxes and their contribution to ecosystem carbon exchange in a pine forest: an assessment based on eddy covariance measurements and an integrated model.

We used a combination of eddy flux, canopy, soil and environmental measurements with an integrated biophysical model to analyze the seasonality of component carbon (C) fluxes and their contribution to ecosystem C exchange in a 50-year-old Scots pine forest (Pinus sylvestris L.) in eastern Finland (62 degrees 47' N, 30 degrees 58' E) over three climatically contrasting years (2000-2002). Eddy flux measurements showed that the growing Scots pine forest was a sink for CO2, with annual net C uptakes of 131, 210 and 258 g C m-2> year-1 in 2000, 2001 and 2002, respectively. The integrated process model reproduced the annual course of daily C flux above the forest canopy as measured by the eddy covariance method once the site-specific component parameters were estimated. The model explained 72, 66 and 68% of the variation in daily net C flux in 2000, 2001 and 2002, respectively. Modeled annual C loss by respiration was 565, 629 and 640 g C m-2 year-1, accounting for 77, 77 and 65% of annual gross C uptake, respectively. Carbon fluxes from the forest floor were the dominant contributors to forest ecosystem respiration, with the fractions of annual respiration from the forest floor, foliage and wood being 46-62, 27-44 and 9-10%, respectively. The wide range in daily net C uptake during the growing season was largely attributable to day-to-day fluctuations in incident quantum irradiance. During just a few days in early spring and late autumn, ecosystem net C exchange varied between source and sink as a result of large daily changes in temperature. The forest showed a greater reduction in gross C uptake by photosynthesis than in C loss by respiration during the dry summer of 2000, indicating that interannual variability in ecosystem net C uptake at this site was modified mostly by summer rainfall and vapor pressure deficit.

Carbon↗
PubMed2004

Spatial and temporal variations in leaf area index, specific leaf area and leaf nitrogen of two co-occurring savanna tree species.

Foliage growth, mass- and area-based leaf nitrogen concentrations (Nm and N a) and specific leaf area (SLA) were surveyed during a complete vegetation cycle for two co-occurring savanna tree species: Crossopteryx febrifuga (Afzel. ex G. Don) Benth. and Cussonia arborea A. Rich. The study was conducted in the natural reserve of Lamto, Ivory Coast, on isolated and clumped trees. Leaf flush occurred before the beginning of the rainy season. Maximum leaf area index (LAI), computed on a projected canopy basis for individual trees, was similar (mean of about 4) for both species. Seasonal courses of the ratio of actual to maximum LAI were similar for individuals of the same species, but differed between species. For C. febrifuga, clumped trees reached their maximum LAI before isolated trees. The LAI of C. arborea trees did not differ between clumped and isolated individuals, but maximum LAI was reached about 2 months later than for C. febrifuga. Leaf fall was associated with decreasing soil water content for C. arborea. For C. febrifuga, leaf fall started before the end of the rainy period and was independent of changes in soil water content. These features lead to a partial niche separation in time for light resource acquisition between the two species. Although Nm, N a and SLA decreased with time, SLA and N a decreased later in the vegetation cycle for C. arborea than for C. febrifuga. For both species, N a decreased and SLA increased with decreasing leaf irradiance within the canopy, although effects of light on leaf characteristics did not differ between isolated and clumped trees. Given relationships between N a and photosynthetic capacities previously reported for these species, our results show that C. arborea exhibits higher photosynthetic capacity than C. febrifuga during most of the vegetation cycle and at all irradiances.

Araliaceae↗
PubMed2004

Stomatal sensitivity to vapor pressure deficit and its relationship to hydraulic conductance in Pinus palustris.

We studied the response of stomatal conductance at leaf (gS) and canopy (GS) scales to increasing vapor pressure deficit (D) in mature Pinus palustris Mill. (longleaf pine) growing in a sandhill habitat in the coastal plain of the southeastern USA. Specifically, we determined if variation in the stomatal response to D was related to variation in hydraulic conductance along the soil-to-leaf pathway (KL) over the course of a growing season. Reductions in KL were associated with a severe growing season drought that significantly reduced soil water content (theta) in the upper 90-cm soil profile. Although KL recovered partially following the drought, it never reached pre-drought values. Stomatal sensitivity to D was well correlated with maximum gS at low D at both leaf and canopy scales, and KL appeared to influence this response by controlling maximum gS. Our results are consistent with the hypothesis that stomatal response to D occurs to regulate minimum leaf water potential, and that the sensitivity of this response is related to changes in whole-plant hydraulics.

Pinus↗
PubMed2005

Assessing forest structure and function from spectral transmittance measurements: a case study in a Mediterranean holm oak forest.

Annual changes in structural attributes and seasonal dynamics in water content, photosynthetic rate and light-use efficiency (LUE) were assessed by spectral transmittance for 4 years (1999-2003) in six stands of a Mediterranean holm oak forest. Green biomass, total biomass and leaf area index (LAI) were determined. In 1999, seasonal dynamics of net carbon dioxide (CO2) exchange and water content were measured. We recorded photosynthetically active radiation (PAR) transmittance and hyperspectral transmittance in the 400-1100 nm region and derived reflectance-based vegetation indices. Transmittance over the PAR region derived from either ceptometer or spectroradiometer measurements (PART and TPAR, respectively) was related to green and total biomass. Both PART and TPAR were also related to LAI (r=0.79 and r=0.70, respectively, P <0.001) and were appropriate for comparison among stands, whereas subtle changes in LAI within a stand were better assessed by the transmittance amplitude in the red edge region (TRE) (within a stand, r=0.77-0.99, P <0.001). Spectral transmittance-based indices successfully captured physiological processes that occurred on temporal (seasonal) and spatial scales. The transmittance-based water index (TWI) was related to both foliage and canopy water content (r=0.69, P <0.001). Estimates of foliage and canopy water content improved in dense (closed) stands (r=0.84 and r=0.87, respectively, P <0.001) compared with low-density stands. Under non-drought conditions, transmittance-based photochemical reflectance index (TPRI) was related to LUE (r=0.58, P <0.05) and net CO2 exchange (r=0.72, P <0.01), and the combined TPAR x TPRI index greatly improved these relationships (r=0.93 and r=0.84, respectively, P <0.01), indicating that both structural and physiological adjustments modified CO2 fixation capacity in these forest stands. Our novel approach to the study of transmitted radiation provides a tool for estimating structural and functional variables such as LAI, LUE and water content, which are key determinants of terrestrial productivity.

Biomass↗
PubMed2005

Production physiology of three fast-growing hardwood species along a soil resource gradient.

We determined how specific leaf area (SLA), specific leaf nitrogen (SLN), leaf area index (LAI), light-saturated photosynthesis (Amax) and aboveground net primary productivity (ANPP) of three commercially important hardwood species, eastern cottonwood (Populus deltoides Bartr.), American sycamore (Platanus occidentalis L.) and cherrybark oak (Quercus falcata var.pagodafolia Ell.), vary across a soil resource gradient. Five treatments were applied in a randomized block design (control, irrigation only (IRR), and irrigation plus fertilization with 56, 112 or 224 kg N ha-1 year-1 (N56, N112 and N224)) with four replications per species. When trees were 6 years old, Amax, SLA, SLN, LAI and ANPP were quantified during peak leaf production within a single growing season. In all species, Amax for sun leaves was significantly higher than for shade leaves (34, 32 and 29 micromol m2 s-1 versus 27, 23 and 23 micromol m2 s-1 for cottonwood, cherrybark oak and sycamore sun and shade leaves, respectively) and tended to plateau in the N112 treatment. The SLA was significantly lower in sun than in shade leaves and reached a plateau in IRR-treated cottonwood and sycamore, and in N56-treated oak. Values of SLN peaked in the N122 treatment for cottonwood sun leaves (1.73 g N m2) and in the N56 treatment for sycamore and oak (1.54 and 1.90 g N m2, respectively). In sun and shade leaves of all species, Amax increased with increasing SLN. Cherrybark oak LAI reached a plateau across the resource gradient in the N56 treatment, whereas cottonwood and sycamore LAI reached a plateau in the IRR treatment. All species exhibited significant curvilinear relationships between canopy Amax and ANPP. These findings indicate that nutrients and water regulate leaf-level traits such as SLA and SLN, which in turn influence LAI and canopy photosynthesis, thereby affecting ANPP at the tree and stand levels.

Biomass↗
PubMed2005

Elevated CO2 concentration, fertilization and their interaction: growth stimulation in a short-rotation poplar coppice (EUROFACE).

We investigated the individual and combined effects of elevated CO2 concentration and fertilization on aboveground growth of three poplar species (Populus alba L. Clone 2AS-11, P. nigra L. Clone Jean Pourtet and P. x euramericana Clone I-214) growing in a short-rotation coppice culture for two growing seasons after coppicing. Free-air carbon dioxide enrichment (FACE) stimulated the number of shoots per stool, leaf area index measured with a fish-eye-type plant canopy analyzer (LAIoptical), and annual leaf production, but did not affect dominant shoot height or canopy productivity index. Comparison of LAIoptical with LAI estimates from litter collections and from allometric relationships showed considerable differences. The increase in biomass in response to FACE was caused by an initial stimulation of absolute and relative growth rates, which disappeared after the first growing season following coppicing. An ontogenetic decline in growth in the FACE treatment, together with strong competition inside the dense plantation, may have caused this decrease. Fertilization did not influence aboveground growth, although some FACE responses were more pronounced in fertilized trees. A species effect was observed for most parameters.

Biomass↗
PubMed2005

Relationships between net photosynthesis and foliar nitrogen concentrations in a loblolly pine forest ecosystem grown in elevated atmospheric carbon dioxide.

We examined the effects of elevated carbon dioxide concentration ([CO2]) on the relationship between light-saturated net photosynthesis (A(sat)) and area-based foliar nitrogen (N) concentration (N(a)) in the canopy of the Duke Forest FACE experiment. Measurements of A(sat) and N(a) were made on two tree species growing in the forest overstory and four tree species growing in the forest understory, in ambient and elevated [CO2] FACE rings, during early and late summer of 1999, 2001 and 2002, corresponding to years three, five and six of CO2 treatment. When measured at the growth [CO2], net photosynthetic rates of each species examined in the forest overstory and understory were stimulated by elevated [CO2] at each measurement date. We found no effect of elevated [CO2] on N(a) in any of the species. The slope of the A(sat)-N relationship was 81% greater in elevated [CO2] than in ambient [CO2] when averaged across all sample dates, reflecting a differential CO2 effect on photosynthesis at the top and bottom of the canopy. We compared A(sat)-N relationships in trees grown in ambient and elevated [CO2] at two common CO2 concentrations, during late summer 2001 and both early and late 2002, to determine if the stimulatory effect of elevated [CO2] on photosynthesis diminishes over time. At all three sample times, neither the slopes nor the y-intercepts of the A(sat)-N relationships of trees grown in ambient or elevated [CO2] differed when measured at common CO2 concentrations, indicating that the responses of photosynthesis to long-term elevated [CO2] did not differ from the responses to a short-term increase in [CO2]. This finding, together with the observation that N(a) was unaffected by growth in elevated [CO2], indicates that these overstory and understory trees growing at the Duke Forest FACE experiment continue to show a strong stimulation of photosynthesis by elevated [CO2].

Atmosphere↗
PubMed2005

Reimplementation of the Biome-BGC model to simulate successional change.

Biogeochemical process models are increasingly employed to simulate current and future forest dynamics, but most simulate only a single canopy type. This limitation means that mixed stands, canopy succession and understory dynamics cannot be modeled, severe handicaps in many forests. The goals of this study were to develop a version of Biome-BGC that supported multiple, interacting vegetation types, and to assess its performance and limitations by comparing modeled results to published data from a 150-year boreal black spruce (Picea mariana (Mill.) BSP) chronosequence in northern Manitoba, Canada. Model data structures and logic were modified to support an arbitrary number of interacting vegetation types; an explicit height calculation was necessary to prioritize radiation and precipitation interception. Two vegetation types, evergreen needle-leaf and deciduous broadleaf, were modeled based on site-specific meteorological and physiological data. The new version of Biome-BGC reliably simulated observed changes in leaf area, net primary production and carbon stocks, and should be useful for modeling the dynamics of mixed-species stands and ecological succession. We discuss the strengths and limitations of Biome-BGC for this application, and note areas in which further work is necessary for reliable simulation of boreal biogeochemical cycling at a landscape scale.

Atmosphere↗
PubMed2005

Vertical, horizontal and azimuthal variations in leaf photosynthetic characteristics within a Fagus crenata crown in relation to light acclimation.

An understanding of spatial variations in gas exchange parameters in relation to the light environment is crucial for modeling canopy photosynthesis. We measured vertical, horizontal and azimuthal (north and south) variations in photosynthetic capacity (i.e., the maximum rate of carboxylation: Vcmax), nitrogen content (N), leaf mass per area (LMA) and chlorophyll content (Chl) in relation to relative photosynthetic photon flux (rPPF) within a Fagus crenata Blume crown. The horizontal gradient of rPPF was similar in magnitude to the vertical gradient of rPPF from the upper to the lower crown. The rPPF in the north quadrant of the crown was slightly lower than in the south quadrant. Nitrogen content per area (Narea), LMA and Vcmax were strictly proportional to rPPF, irrespective of the vertical direction, horizontal direction and crown azimuth, whereas nitrogen content per dry mass, Chl per area and photosynthetic capacity per dry mass (Vm) were fairly constant. Statistical analyses separating vertical trends from horizontal and azimuthal trends indicated that, although horizontal and vertical light acclimation of leaf properties were similar, there were two significant azimuthal variations: (1) Vcmax was lower in north-facing leaves than in south-facing leaves for a given Narea, indicating low photosynthetic nitrogen-use efficiency (PNUE) of north-facing leaves; and (2) Vcmax was lower in north-facing leaves than in south-facing leaves for a given LMA, indicating low Vm of the north-facing leaves. With respect to the low PNUE of the north-facing leaves, there were no significant azimuthal variations in leaf CO2 conductance from the stomata to the carboxylation site. Biochemical analysis indicated that azimuthal variations in nitrogen allocation to ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) and in nitrogen allocation between carboxylation (Rubisco and other Calvin cycle enzymes) and light harvesting machinery (Chl pigment-protein complexes) were not the main contributor to the difference in PNUE between north- and south-facing leaves. Lower specific activity of Rubisco may be responsible for the low PNUE of the north-facing leaves. Anatomical analysis indicated that not only high leaf density, which is compatible with a greater fraction of non-photosynthetic tissue, but also thick photosynthetic tissue contributed to the low Vm in the north-facing leaves. These azimuthal variations may need to be considered when modeling canopy photosynthesis based on the Narea-Vcmax or LMA-Vcmax relationship.

Acclimatization↗
PubMed2005

Photosynthetic acclimation to light changes in tropical monsoon forest woody species differing in adult stature.

We studied morphological and physiological leaf and whole-plant features of seedlings of six late-successional woody species common in the Xishuangbanna lowland rain forest in southwest China. Study species differed in adult stature and shade tolerance and included the shrubs Lasianthus attenuatus Jack and Lasianthus hookeri C.B. Clarke ex Hook. f.; the sub-canopy species Barringtonia macrostachya (Jack) Kurz and Linociera insignis C.B. Clarke; the canopy tree Pometia tomentosa (Blume) Teijsm. & Binn.; and the emergent species Shorea chinensis (Wang Hsie) H. Zhu. After 1 year of growth in low light (4.5% full sun), seedlings were transferred to high light (24.5% full sun) to investigate acclimation responses of existing leaves to forest gap opening and to determine whether seedling capacity for acclimation is a limiting factor in its natural regeneration. Leaves of the shrub species are shade-adapted, as indicated by their low photosynthetic capacity, efficiency in using sunflecks, low stomatal density, low Chl a/b ratio and high spongy/palisade mesophyll ratio. The shrub species utilized sunflecks efficiently because they had a short photosynthetic induction time and low induction loss. In all species, transfer of seedlings to high light resulted in a substantial initial reduction in the dark-adapted quantum yield of photosystem II (variable chlorophyll fluorescence/maximum chlorophyll fluorescence; Fv/Fm) at midday. Predawn Fv/Fm of the taller species did not change greatly, but predawn Fv/Fm of the shrub species decreased significantly without complete recovery within 25 days of transfer to high light, indicating chronic photoinhibition and damage to the previously shade-adapted leaves. Maximum net photosynthetic rate and dark respiration of the four taller species increased considerably after transfer to high light, but not in the shrub species. Similar trends were observed for the number of newly formed leaves and relative height growth rate. We conclude that the shrubs L. hookeri and L. attenuatus have limited potential for developmental and physiological acclimation to high light, which explains their absence from forest gaps. Compared with the shrub species, the taller tree species, which are more likely to experience high light during their life span, showed a greater potential for light acclimation. Physiological differences among the four tree species were not consistent with differences in adult stature.

Acclimatization↗
PubMed2006

Testing the unifying theory of ozone sensitivity with mature trees of Fagus sylvatica and Picea abies.

The broad range in plant responses to chronic O(3) exposure compels a search for integrative, underlying principles. One such approach is the unifying theory proposed by Reich (1987), which combines the O(3) response of contrasting physiognomic classes of plants on the basis of their intrinsic leaf diffusive conductance and, hence, capacity for O(3) uptake. Physiognomic classes differ in the proportional decline in photosynthesis and growth when compared on the basis of cumulative O(3) exposure per unit time, but converge when compared on the basis of O(3) uptake per unit time or cumulative O(3) uptake over the entire lifetime of the leaf. The theory is based on observations on a large number of contrasting plant species, relying primarily on studies of juvenile trees subjected to short-term O(3) exposure. To test the applicability of the unifying theory to mature trees, broadleaf deciduous European beech (Fagus sylvatica L.) and the evergreen conifer Norway spruce (Picea abies (L.) Karst.) in a mature mixed stand were exposed to either ambient air (control) or air with twice the ambient O(3) concentration delivered into the canopy by means of a free-air fumigation system. We accounted for differences in growing season length, leaf longevity and O(3)-related effects on leaf diffusive conductance in determining total O(3) uptake over the lifetime of the leaf. On this basis, Norway spruce needles required 5 years to take up as much O(3) as did beech leaves in one growing season. The core of the unifying theory on O(3) sensitivity was substantiated in relation to O(3) exposure and uptake. However, contrary to the unifying theory, which was formulated on the basis of results with juvenile trees, the O(3) response of mature trees in a natural stand was more complex. The increased complexity was attributed to additional environmental stressors, stress compensation at the whole-tree level, and differential O(3) sensitivities of leaves according to age class and position within the canopy. Contrary to the theory, photosynthesis was no less sensitive to O(3) in Norway spruce than that of beech, and was reduced in the twice-ambient O(3) regime in the first year of exposure.

Fagus↗
PubMed2006

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.

Australia↗
PubMed2007

Changes in net ecosystem productivity with forest age following clearcutting of a coastal Douglas-fir forest: testing a mathematical model with eddy covariance measurements along a forest chronosequence.

We hypothesized that changes in net ecosystem productivity (NEP) during aging of coastal Douglas-fir (Pseudotsuga menziesii Mirb. Franco) stands could be explained by (1) changing nutrient uptake caused by different time scales for decomposition of fine, non-woody and coarse woody litter left after harvesting, (2) declines in canopy water status with lengthening of the water uptake pathway during bole and branch growth, and (3) increases in the ratio of autotrophic respiration (R (a)) to gross primary productivity (GPP) with phytomass accumulation. These hypotheses were implemented and tested in the mathematical model ecosys against eddy covariance (EC) measurements of forest CO(2) and energy exchange in a post-clearcut Douglas-fir chronosequence. Hypothesis 1 explained how (a) an initial rise in GPP observed during the first 3 years after clearcutting could be caused by nutrient mineralization from rapid decomposition of fine, non-woody litter with lower C:N ratios (assart effect), (b) a slower rise in GPP during the next 20 years could be caused by immobilization during later decomposition of coarse woody litter, and (c) a rapid rise in GPP between 20 and 40 years after clearcutting could be caused by nutrient mineralization with further decomposition of coarse woody litter and of its decomposition products. During periods (a) and (b), heterotrophic respiration (R (h)) from decomposition of fine and coarse litter greatly exceeded net primary productivity (NPP = GPP - R (a)) so that Douglas-fir stands were large sources of CO(2). During period (c), NPP exceeded R (h) so that these stands became large sinks for CO(2). Hypothesis 2 explained how declines in NPP during later growth in period (c) could be caused by lower hydraulic conductances in taller trees that would force lower canopy water potentials and hence greater sensitivity of stomatal conductances and CO(2) uptake to vapor pressure deficits. Enhanced sensitivity to vapor pressure deficits was also apparent in the EC measurements over the post-clearcut chronosequence. Hypothesis 3 did not contribute to the explanation of forest age effects on NEP.

Abies↗