Leaf distribution in large trees and stands of the floodplain forest in southern Moravia

Vertical distributions of leaf dry mass (M(d)) and leaf area (A(f)) were related to relative irradiance (I(r); I(r) above the stand = 1) in closed-canopy, old-growth stands of the floodplain forest in southern Moravia composed largely of Quercus, Fraxinus and Tilia species. Foliage area and mass at any given canopy height were converted to solar equivalent leaf area (A(s)) and mass (M(s)) by multiplying actual values at a given level in the canopy by the relative irradiance at that position. Stand leaf area index (LAI) was 5 (7 including shrub and herb layer), and solar equivalent parameters reached about 25% of that amount. In all species, vertical profiles of both relative irradiance and leaf dry mass to area ratio (LMA) were sigmoidal and the two variables were linearly related. The dominant, upper canopy species had a larger proportion of solar equivalent foliage than suppressed understory species. For individual trees of all species, the upper canopy had a larger proportion of solar equivalent foliage than the lower canopy. Light compensation points at both the leaf and whole-tree level were defined according to leaf or tree position, size and structure. I conclude that optimization of A(s) for forest stands may be used as a basis for determining thinning schedules and evaluating tree survival after damage to tree crowns by various factors.     

Variations in relative stomatal and biochemical limitations to photosynthesis in a young blackbutt (Eucalyptus pilularis) plantation subjected to different weed control regimes

Foliar gas exchange and carbon (delta(13)C) and oxygen (delta(18)O) isotope ratios were measured in a young blackbutt (Eucalyptus pilularis Sm.) plantation subjected to four weed control treatments defined by the width of the weed-free strip maintained for the first 12 months after planting. Treatments were: 2-m-wide weed-free strip (50% of plot area, 2.0MWC), 1.5-m-wide weed-free strip (37.5% of plot area, 1.5MWC), 1-m-wide weed-free strip (25% of plot area, 1.0MWC) and no weed control (NWC). Our objectives were to determine (1) if decreasing the width of the weed control strip (decreasing herbicide use) affected growth and leaf photosynthesis of the plantation, and (2) the effects of the weed control regimes on variations in relative stomatal and biochemical limitations to photosynthesis. Trees in the 1.0MWC treatment had lower foliar light-saturated photosynthetic rate (A(sat)) than trees in the 2.0MWC treatment. An increase in metabolic limitation was responsible for the decrease in A(sat) in the 1.0MWC trees, which was also partly confirmed by the isotopic data. Compared with trees in the 1.0MWC, 1.5MWC and 2.0MWC treatments, A(sat) of NWC trees was significantly lower, a difference that was attributable mainly to stomatal limitation and to a lesser extent to biochemical limitation. The results support the conclusion that different weed control regimes cause differences in relative stomatal and biochemical limitations to plantation photosynthesis. This report contributes to a growing body of literature on competition for soil resources between trees and weeds. Our results highlight the usefulness of the stable isotopic method in supporting analysis of the response of net photosynthesis to varying intercellular CO(2) concentration for determining the relative stomatal and non-stomatal limitations to photosynthesis under experimental conditions in the field.     

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.     

Parameterization and testing of a biochemically based photosynthesis model for walnut (Juglans regia) trees and seedlings

The biochemically based leaf photosynthesis model proposed by Farquhar et al. (1980) and the stomatal conductance model proposed by Jarvis (1976) were parameterized for walnut. Responses of photosynthesis to CO(2) and irradiance were used to determine the key parameters of the photosynthesis model. Concurrently, stomatal conductance responses to leaf irradiance (Q), leaf temperature (T(l)), water vapor pressure deficit at the leaf surface (D), and air CO(2) concentration at the leaf surface (C(s)) were used to parameterize the stomatal conductance model. To test the generality of the model parameters, measurements were made on leaves from a 20-year-old tree growing in the field, and from sunlit and shaded greenhouse-grown seedlings. The three key parameters of the photosynthesis model (maximum carboxylation rate V(cmax), electron transport capacity J(max), and dark respiration rate R(d)) and the key parameter of the conductance model (reference stomatal conductance, g(sref)) were linearly correlated with the amount of leaf nitrogen per unit leaf area. Unique relationships could be used to describe nitrogen effects on these parameters for leaves from both the tree and the seedlings. Our data allowed separation of the effects of increasing total photosynthetic apparatus per unit leaf area from the effects of partitioning nitrogen among different pools of this apparatus for foliage acclimation to leaf irradiance. Strong correlations were found between stomatal conductance g(s) and Q, D and C(s), whereas the relationship between g(s) and T(l) was weak. Based on these parameterizations, the model adequately predicted leaf photosynthesis and stomatal conductance when tested with an independent set of data obtained for the tree and seedlings. Total light-driven electron flows derived from chlorophyll fluorescence data obtained at different leaf temperatures were consistent with values computed by the model. The model was also tested with branch bag data acquired from a three-year-old potted walnut tree. Despite a relatively large variance between observed and simulated values, the model predicted stomatal conductance and photosynthesis reasonably well at the branch scale. The results indicate that the photosynthesis-conductance model developed here is robust and can be applied to walnut trees and seedlings under various environmental conditions where water is non-limiting.     

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].     

Light environment alters ozone uptake per net photosynthetic rate in black cherry trees

Foliar ozone uptake rates of different-sized black cherry (Prunus serotina Ehrh.) trees were compared within a deciduous forest and adjacent openings in north-central Pennsylvania during one growing season. Study trees included open-grown seedlings and saplings, forest understory seedlings and saplings, and sunlit and shaded portions of mature canopy tree crowns. Instantaneous ozone uptake rates were highest in high-light environments primarily because of higher stomatal conductances. Low ozone uptake rates of seedlings and saplings in the forest understory could be attributed partially to lower average ambient ozone concentrations compared to the canopy and open environments. Among the tree size and light combinations tested, ozone uptake rates were highest in open-grown seedlings and lowest in forest-grown seedlings. Despite lower ozone uptake rates of foliage in shaded environments, ozone uptake per net photosynthesis of foliage in shaded environments was significantly higher than that of foliage in sunlit environments because of weaker coupling between net photosynthesis and stomatal conductance in shaded environments. The potential for greater ozone injury in shaded environments as a result of greater ozone uptake per net photosynthesis is consistent with previous reports of greater ozone injury in shaded foliage than in sunlit foliage.     

Photosynthetic capacity in relation to nitrogen in the canopy of a Quercus robur, Fraxinus angustifolia and Tilia cordata flood plain forest

We measured gas exchange and various leaf parameters of ash (Fraxinus angustifolia Vahl.) and oak (Quercus robur L.) in the high canopy and of lime (Tilia cordata Mill.) in the lower canopy of a planted, 120-year-old floodplain forest in southern Moravia, Czech Republic. The high-canopy leaves of F. angustifolia and Q. robur had nitrogen concentrations on a leaf area basis (N(area)) that were twice those of low-canopy leaves of T. cordata. Upper-canopy leaves of F. angustifolia had a photosynthetic rate at light saturation (A(max)) of about 16 micromol CO2 m(-2) s(-1), whereas A(max) of the upper-canopy foliage of Q. robur achieved only about two thirds of this value. Contrary to previous investigations of photosynthetic performance in monospecific stands, leaves of the uppermost branches of T. cordata at 15-m height had the highest A(max) and transpiration rate among the species studied. Water-use efficiency (WUE) was low in T. cordata at 15-m canopy height, whereas WUE was significantly higher for Q. robur leaves at 27-m height than for the other species. Leaves of T. cordata at 15-m height showed the strongest relationship between A(max) and N(area) (R2 = 0.90) followed by F. angustifolia (R2 = 0.69). The strong correlation between photosynthesis and nitrogen concentration in T. cordata at 15 m, together with the steep regression slope for the A(max):N(area) relationship, indicated that nitrogen allocation to the photosynthetic apparatus resulted in high nitrogen-use efficiency of light-saturated photosynthesis (PNUE). Despite differences in PNUE among species, PNUE was fairly constant for leaves sampled from the same canopy position, suggesting that single-leaf parameters are matched to optimize PNUE for prevailing light conditions. High PNUE in T. cordata at 15 m partially compensated for the species' subordinate position in the canopy, and may be an important mechanism for its coexistence in highly structured vegetation.     

Why does leaf nitrogen decline within tree canopies less rapidly than light? An explanation from optimization subject to a lower bound on leaf mass per area

A long-established theoretical result states that, for a given total canopy nitrogen (N) content, canopy photosynthesis is maximized when the within-canopy gradient in leaf N per unit area (N(a)) is equal to the light gradient. However, it is widely observed that N(a) declines less rapidly than light in real plant canopies. Here we show that this general observation can be explained by optimal leaf acclimation to light subject to a lower-bound constraint on the leaf mass per area (m(a)). Using a simple model of the carbon-nitrogen (C-N) balance of trees with a steady-state canopy, we implement this constraint within the framework of the MAXX optimization hypothesis that maximizes net canopy C export. Virtually all canopy traits predicted by MAXX (leaf N gradient, leaf N concentration, leaf photosynthetic capacity, canopy N content, leaf-area index) are in close agreement with the values observed in a mature stand of Norway spruce trees (Picea abies L. Karst.). An alternative upper-bound constraint on leaf photosynthetic capacity (A(sat)) does not reproduce the canopy traits of this stand. MAXX subject to a lower bound on m(a) is also qualitatively consistent with co-variations in leaf N gradient, m(a) and A(sat) observed across a range of temperate and tropical tree species. Our study highlights the key role of constraints in optimization models of plant function.     

Seasonal and yearly variations in light use and nitrogen use by seedlings of four deciduous broad-leaved tree species invading larch plantations

Several deciduous broad-leaved tree species, differing in leaf phenology, invade larch (Larix kaempferii (Lamb.) Carrière) plantations in Japan. The understory light environment of larch forests changes drastically between the leafy and leafless periods. To determine how the invading seedlings exploit the changing light environment, and if phenological differences reflect the light- and nitrogen-use traits of the seedlings, we measured leaf phenology, seasonal changes in light-saturated photosynthetic rate (P(sat)), leaf nitrogen (N) content (N(area)), chlorophyll/nitrogen ratio (Chl/N), specific leaf area (SLA) and N remobilization rate (NRMR) over 3 years. The mid-successional or gap-phase species, Magnolia hypoleuca Siebold & Zucc., had a short leafy period and high P(sat) and NRMR. In contrast, two late-successional tree species, Prunus ssiori Friedr. Schmidt, which undergoes leaf flush before larch, and Carpinus cordata Blume, which maintains green leaves until frost, both had low P(sat) and NRMR but exploited the opportunity for growth during the period when the larch canopy trees were leafless. Quercus mongolica Fisch. ex Ledeb. var. crispula (Blume) Ohashi, a mid-late-successional species that underwent leaf flush at the same time as the overstory larch, had values of photosynthetic parameters between those of the gap-phase and late-successional species. Among species, M. hypoleuca and Q. mongolica had higher photosynthetic rates and photosynthetic N-use efficiencies. In all species, the relationship between N(area) and P(sat) showed species-specific yearly fluctuations; however, there was no yearly fluctuation in the relationship between N(area) and P(sat) at CO2 saturation. Yearly fluctuations in the N(area)-P(sat) relationship appeared to be induced by changes in SLA and N-use characteristics, which in turn are affected by climatic variations.     

Changes in photosynthesis and water status of developing leaves of Brachystegia spiciformis Benth

Changes in net carbon assimilation and water status were studied during leaf development in the deciduous, tropical species Brachystegia spiciformis Benth. In this upland savanna African tree, bud-burst and leaf development occur approximately two months before the rainy season. The newly formed leaves synthesize anthocyanin until the fully expanded leaves of the whole canopy are red. This foliage is referred to as "spring flush" foliage. Subsequently, the anthocyanins are metabolized and the pre-rain leaves become green. Carbon dioxide assimilation exhibited a bimodal diurnal pattern and was similar for pre-rain green leaves and fully expanded flushing leaves, although pre-rain green leaves showed a net uptake of carbon throughout the daylight period, whereas flushing leaves exhibited only brief periods of net photosynthesis in the morning and early afternoon. Measurements of leaf water potential and relative water content showed a diurnal pattern with considerable variation throughout the day. Leaf water potential and relative water content values decreased soon after sunrise reaching a minimum at a time corresponding to the afternoon peak in CO(2) assimilation. Stomatal conductance was closely related to transpiration rate in both flushing and pre-rain green leaves, although flushing leaves had lower stomatal conductances than pre-rain green leaves. Pre-rain green leaves exhibited a compensation irradiance of approximately 180 micro mol m(-2) s(-1), whereas flushing leaves had positive net photosynthesis only at PPFDs greater than 300 micro mol m(-2) s(-1). Rate of photosynthesis (expressed per leaf area or chlorophyll unit) increased as anthocyanin concentration decreased, although the photosynthetic rate continued to increase long after the leaf anthocyanins had been degraded to low, visually undetectable amounts. Post-rain green leaves had chlorophyll concentrations, transpiration rates and stomatal conductances similar to those of pre-rain green leaves; however, photosynthetic rates in post-rain leaves were more than three times higher. Thus, during the early stages of the spring flush, carbon asimilation rates of the flushing leaves were inversely related to leaf anthocyanin concentrations. In pre-rain green leaves, photosynthesis was limited by other non-stomatal factors.     

Constraints on light interception efficiency due to shoot architecture in broad-leaved Nothofagus species

Shoot architecture may significantly alter mean quantum flux on foliage and thus, photosynthetic productivity. There is currently only limited information about plastic alterations in shoot structure caused by within-canopy variation in mean integrated irradiance (Q(int)) in broad-leaved trees. We studied leaf and shoot structure, and nitrogen and carbon content in late-successional, widely distributed, temperate, broad-leaved Nothofagus taxa to determine the architectural controls on light harvesting and photosynthetic performance. Nothofagus fusca (Hook. f.) Oersted has larger leaves and less densely leaved shoots than the N. solandri varieties. Nothofagus solandri var. solandri (Hook. f.) Oersted is characterized by rounder leaves that potentially have a larger overlap than the ovate-triangular leaves of N. solandri var. cliffortioides (Hook. f.) Poole. Leaf dry mass (M(A)) and nitrogen content (N(A)) per unit area increased with increasing Q(int) in all species, demonstrating enhanced investment of photosynthetic biomass in high light. Although M(A) differed between species at a common irradiance, there was a uniform relationship between N(A) and Q(int) across species. Leaf carbon content per dry mass and leaf dry mass to fresh mass ratio also scaled positively with irradiance, suggesting greater structural investments in high light. In all species, shoots became more horizontal and flatter at lower Q(int), implying an enhanced use efficiency of direct irradiance in natural leaf positions. In contrast, irradiance effects on leaf aggregation varied among species. Across the data, leaf overlap or leaf area density was often greater at lower irradiances, possibly as a result of limited carbon availability for shoot axis extension growth. In N. fusca, leaves of which were more aggregated in high light, the shoot silhouette to total leaf area ratio (S(S)) declined strongly with increasing irradiance, demonstrating a lower light harvesting efficiency at high Q(int). This effect was only moderate in N. solandri var. cliffortioides and S(S) was independent of Q(int) in N. solandri var. solandri. Although the efficiency of light interception at high irradiances was lowest in N. fusca, this species had the greatest nitrogen content per unit shoot silhouette area (2N(A)/S(S)), indicating superior shoot-level photosynthetic potential. These data collectively demonstrate that shoot architecture significantly affects light interception and photosynthesis in broad-leaved trees, and that structural carbon limitations may constrain leaf light harvesting efficiency at low irradiance.     

Optimality and nitrogen allocation in a tree canopy

Physical and functional properties of foliage were measured at a variety of microsites in a broad-leaved Nothofagus fusca (Hook. f.) ?rst. canopy. The light climate of the foliage at these sites was monitored for 39 days in the late spring and early summer with in situ sensors. Foliage nitrogen content (N), mean leaf angle, and gas exchange characteristics were all correlated with the amount of light reaching the microsites during foliage development. Foliage N content on a leaf area basis ranged between ~1 and 2.5 g N m(-2) and was highest at the brightest sites. Light-saturated photosynthetic rates ranged between ~4 and 9 micro mol m(-2) s(-1), increasing from the darkest to brightest sites. A biochemical model of photosynthesis was fitted to foliage characteristics at the different microsites and used to integrate foliage assimilation among the sites over 39 days. The actual arrangement of foliage physiological characteristics in the observed microsites led to higher total canopy rates of net assimilation than > 99% of the combinations of observed foliage characteristics randomly assigned to the observed microsites. Additional simulations first related the maximum rates of electron transport (J(max)), ribulose bisphosphate turnover (V(c,max)), and dark respiration (R(d)) of Nothofagus fusca foliage to nitrogen content and then allowed foliage N (and consequently leaf gas exchange characteristics) to vary across the canopy. The observed N allocation pattern results in greater total canopy assimilation than uniform or > 99% of the simulations with random distributions of N among the microsites (constrained so that the total N allocated was equivalent to that observed in the microsites). However, the observed pattern of N allocation places less N in the brightest microsites and results in substantially less total assimilation than a simulated canopy in which N was allocated in an optimal manner where the N distribution is such that the partial derivative of leaf assimilation (A) with respect to leaf nitrogen content, partial differential A/ partial differential N, is constant among microsites. These results suggest that other factors such as wind or herbivory reduce the integrated assimilation of high-N foliage relatively more than lower-N foliage and that a partial differential A/ partial differential N optimality criteria based only on formulations of leaf gas exchange overestimate canopy assimilation.     

Canopy dynamics and aboveground production of five tree species with different leaf longevities

Canopy dynamics and aboveground net primary production (ANPP) were studied in replicated monospecific and dual-species plantations comprised of species with different leaf longevities. In the monospecific plantations, leaf longevity averaged 5, 6, 36, 46 and 66 months for Quercus rubra L., Larix decidua Miller, Pinus strobus L., Pinus resinosa Ait. and Picea abies (L.) Karst., respectively. Specific leaf area, maximum net photosynthesis per unit mass (A/mass), leaf N per unit mass (N(leaf)/mass) and maximum net photosynthesis on a leaf N basis (A/N(leaf)) were inversely correlated to leaf longevity (r(2) = 0.92-0.97, 0.91, 0.88 and 0.80, respectively). Maximum net photosynthesis per unit area (A/area) was not correlated to leaf longevity, whereas leaf N per unit area (N(leaf)/area) was positively correlated to leaf longevity (r(2) = 0.95). For a similar-diameter conifer, species with long-lived foliage supported a greater foliage mass than species with short-lived foliage; however, Quercus rubra did not follow this pattern. At the stand level, total foliage mass ranged from 3.3 to 30.5 Mg ha(-1) and was positively correlated (r(2) = 0.97) to leaf longevity. Leaf area index (LAI) was also positively correlated (r(2) = 0.82) to leaf longevity. Production efficiency (ANPP/LAI) was inversely related to leaf longevity and positively related to A/mass. Aboveground biomass and net primary production differed significantly (P < 0.05) among the five species but were not correlated to leaf longevity, total foliage mass or leaf area. In monospecific plantations, stem NPP for Larix decidua was 17% greater than for Pinus strobus and 14% less than for Picea abies, but in mixed-species plantations stem NPP for Larix decidua was 62 and 85% greater than for Pinus strobus and Picea abies, respectively. Similar aboveground net primary production rates can be attained by tree species with different leaf longevities because of trade-offs resulting from different structural and physiological leaf and canopy characteristics that are correlated to each other and to leaf longevity.     

An estimate of the photosynthetic production of individual trees in a Chamaecyparis obtusa plantation

A Weibull function was used to model the vertical distribution of leaf area of individual trees in a 25-year-old Chamaecyparis obtusa (Siebold & Zucc.) Endl. plantation. The parameter representing the shape of the leaf distribution was independent of tree size. A scale parameter tended to decrease with tree size suggesting a critical minimum height for retention of foliage by trees. On the basis of leaf distribution, the photosynthetic production of individual trees was estimated from the canopy photosynthetic production, which was determined from a model of canopy photosynthesis. The data indicated that the photosynthesis of a tree was proportional to the corresponding tree weight to the power of 1.84. Furthermore, the photosynthetic production varied as the 3/2nd power of total leaf area of the tree. Thus, it was concluded that the photosynthetic production per unit of leaf area, that is, the mean photosynthetic activity of a tree, is proportional to the stem girth at clear length, or the square root of the leaf area of the tree.     

Ontogenetic changes in stomatal and biochemical limitations to photosynthesis of two co-occurring Mediterranean oaks differing in leaf life span

A quantitative analysis was applied to the stomatal and biochemical limitations to light-saturated net photosynthesis under optimal field conditions in mature trees and seedlings of the co-occurring evergreen oak, Quercus ilex L., and the deciduous oak, Q. faginea Lam. Stomatal limitation to photosynthesis, maximal Rubisco activity and electron transport rate were determined from assimilation versus intercellular leaf carbon dioxide concentration response curves of leaves that were subsequently analyzed for nitrogen (N) concentration, mass per unit area, thickness and percent internal air space. In both species, seedlings had a lower leaf mass per unit area, thickness and leaf N concentration than mature trees. The root system of seedlings during their third year after planting was dominated by a taproot. A lower leaf N concentration of seedlings was associated with lower maximal Rubisco activity and electron transport rate and with assimilation rates similar to or lower than those of mature trees, despite the higher stomatal conductances and potential photosynthetic nitrogen-use efficiencies of seedlings. Consequently, stomatal limitation to photosynthesis increased with tree age in both species. In both seedlings and mature trees, a lower assimilation rate in Q. ilex than in Q. faginea was associated with lower stomatal conductance, N allocation to photosynthetic functions, maximal Rubisco activity and electron transport rate, and potential photosynthetic nitrogen-use efficiency but greater leaf thickness and leaf mass per unit area. Tree-age-related changes differed quantitatively between species, and the characteristics of the two species were more similar in seedlings than in mature trees. Despite higher stomatal conductances, seedlings are more N limited than adult trees, which contributes to lower biochemical efficiency.     

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.     

石漠化地区3个树种幼苗在水分胁迫下的光合特性与抗旱性关系

采用盆栽控水方法,研究了翅莢木、滇楸和银荆树3树种当年生实生苗的光合特性与抗旱性的关系.结果表明:随着干旱胁迫强度的增加和时间的延长,3树种的净光合速率(Pn)、蒸腾速率(Tr)和气孔导度(Gs)均呈下降趋势,而胞间CO2浓度则呈上升的趋势;净光合速率日变化曲线逐渐从典型的单峰型转变成双峰型,出现"光合午休"现象.在轻度胁迫和中度胁迫下,净光合速率的降低是由气孔因素和非气孔因素共同引起的:而重度胁迫下,净光合速率的降低主要是非气孔因素引起的.干旱复水后,3个树种的净光合速率均有明显的恢复.利用隶属函数与反隶属函数法对3个树种的抗旱能力进行综合评价,其抗旱能力为:银荆树＞翅莢木＞滇楸.     

Stomatal patchiness in the Mediterranean holm oak (Quercus ilex L.) under water stress in the nursery and in the forest

The evergreen holm oak Quercus ilex L. is the most representative tree in Mediterranean forests. Accurate estimation of the limiting factors of photosynthesis for Q. ilex and the prediction of ecosystem water-use efficiency by mechanistic models can be achieved only by establishing whether this species shows heterogenic stomatal aperture, and, if so, the circumstances in which this occurs. Here, we collected gas-exchange and chlorophyll fluorescence data in Q. ilex leaves from a nursery to measure the effects of stomatal oscillations on PSII quantum yield (Φ(PSII)) under water stress. Stomatal conductance (g(s)) was used as an integrative indicator of the degree of water stress. Images of chlorophyll fluorescence showed heterogeneous Φ(PSII) when g(s) was <50 mmol H(2)O m(-2) s(-1), representative of severe drought and corresponding to a container capacity <45%. Stomatal patchiness was related to a coefficient of variation (CV) of Φ(PSII) values >2.5%. A parallel study in the forest confirmed heterogeneous Φ(PSII) values in leaves in response to declining water availability. Three kinds of Q. ilex individuals were distinguished: those resprouting after a clear-cut (resprouts, R); intact individuals growing in the same clear-cut area as resprouts (controls, C); and intact individuals in a nearby, undisturbed area (forest controls, CF). Patchiness increased in C and CF in response to increasing drought from early May to late July, whereas in R, Φ(PSII) values were maintained as a result of their improved water relations since the pre-existing roots were associated with a smaller aerial biomass. Patchiness was related to a % CV of Φ(PSII) values >4 and associated in the summer with mean g(s) values of 30 mmol H(2)O m(-2) s(-1). Under milder drought in spring, Φ(PSII) patchiness was less strictly related to g(s) variations, pointing to biochemical limitants of photosynthesis. The occurrence of heterogenic photosynthesis caused by patchy stomatal closure in Q. ilex during severe drought should be taken into account in ecosystem modelling in which harsher water stress conditions associated with climate change are predicted.     

粤西地区23个阔叶树种早期生长比较

对南亚热带森林群落恢复标准化示范区23个阔叶树种的早期生长进行了调查与分析。结果表明：23个阔叶树种在本地区有极强的适应性,且在树高、胸径、冠幅生长上均存在着极显著差异,为树种选择提供了依据;米老排、黎蒴栲生长速度最快;红锥、灰木莲、山杜英、锥栗、枫香、樟树生长较快;非洲桃花心木生长速度中等;其他树种生长较慢。     

中山市防火林带树种组成与冠层结构

防火林带的树种组成与冠层结构可影响林带的防火功能。为评价中山市防火林带的树种组成 和冠层结构,对中山市防火林带树种进行了调查,并在防火林带中选取了9 个典型林带类型,建立标准 地,对其树种组成与冠层结构进行了对比研究。结果表明：中山市13 个镇区的生物防火林带共应用了31 个树种,隶属17 科24 属,豆科、山茶科和樟科树种最多,数量最多的有荷木Schima superba、马尾松 Pinus massoniana、台湾相思Acacia confusa、柠檬桉Eucalyptus citriodora、米老排Mytilaria laosensis;以 针叶树为主的针叶林林冠开度最大,达38.07%,林下植物相对茂盛,分别以米老排和杨梅Myrica rubra 为主的阔叶混交林林冠开度最小,分别为12.36% 和13.88%。米老排与杨梅为主要树种的阔叶林林冠开 度比以马尾松为主的针叶林的林冠开度要小,阔叶林的冠层郁闭度较高;自然保护区核心区针阔混交林 的叶面积指数最大,达2.64,显著高于马尾松纯林和马占相思Acacia mangium、荷木和藜蒴Castanopsis fissa 等阔叶混交林,为0.99 和1.65;山脚的荷木林地、荷木与马尾松的混交林和以米老排为主的阔叶混 交林的林下光照较弱,林下植被稀少,冠层结构较为疏散,是较好的防火林带。对现有防火林带的林分 改造,如增加经济林木、果树等阔叶树种,可进一步提高防火林带的防火作用。