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.     

Seasonal variation in the photosynthetic capacity of a willow (Salix cv. Aquatica gigantea) canopy. 2 Comparison of the structure and function of chloroplasts at different levels in the canopy

Development of leaf chloroplast ultrastructure at five levels in a willow (Salix cv. Aquatica gigantea) canopy was followed during one growing season in the field. Changes in chloroplast ultrastructure were compared with the rate of CO(2) uptake of the same leaves. The highest rates of CO(2) uptake were recorded in young leaves exposed to full available sunlight. In these leaves, the area of the grana stacks was less than 20% of the total chloroplast area and the degree of thylakoid stacking was less than 1.5. The chloroplasts of these leaves contained large amounts of starch and small amounts of plastoglobuli. As the canopy grew and the leaves in the lower parts of the canopy became shaded, the structure of the chloroplast thylakoids gradually changed. In leaves at the two lowest levels of the canopy, the degree of stacking at the end of the growing season was close to 2 and correspondingly the rate of CO(2) uptake was low. The areas of grana stacks and plastoglobuli in these chloroplasts increased and were about 30 and 10% of the chloroplast area, respectively, by the end of the growing season. The increase in the degree of thylakoid stacking was caused by increased biosynthesis of grana lamellae, which in general were thinner than the lamellae of young leaves. The length of the stroma lamellae did not change with leaf age. Morphometric measurements showed that the structure of the chloroplasts in leaves 160 cm above ground was dynamic and responsive to environmental conditions so that photosynthetic capacity remained high for 7 weeks despite an increase in leaf shading.     

Seasonal and spatial variations in leaf nitrogen content and resorption in a Quercus serrata canopy

To elucidate the relationships between spatiotemporal changes in leaf nitrogen (N) content and canopy dynamics, changes in leaf N and distribution in the canopy of a 26-year-old deciduous oak (Quercus serrata Thunb. ex. Murray) stand were monitored throughout the developmental sequence from leaf expansion to senescence, by estimating the leaf mass and N concentrations of all the canopy layers. Seasonal changes were observed in leaf N concentration per unit leaf dry mass (N (m)), which peaked after bud burst, declined for two weeks shortly thereafter, and then remained constant for the rest of the growing season for each canopy layer. Leaf N concentration per unit leaf area (N (a)) was higher in the upper layer than in the lower layer throughout the growing season, and was closely correlated with relative irradiance (RI) in the summer when the air temperature was moderately high. The N concentrations of all leaf layers started to decrease in November, and reached their lowest values in late November, whereas LMA scarcely changed throughout the season. The lowest N concentrations did not differ significantly among the canopy layers. Seasonal changes in the relationship between N (a) and RI were detected, indicating that N (a) is optimized temporally as well as spatially. Nitrogen resorption efficiency was highest in the upper canopy layers where larger amounts of N were invested. Based on the estimates of leaf mass and leaf N concentrations of the canopy layers, total leaf N concentration of the whole canopy was estimated to be 84.1 kg ha(-1) in the summer, and 37.3 kg ha(-1) in late November. Therefore, 46.8 kg ha(-1) of leaf N in the canopy (about 56% of the total N) was resorbed just before leaf abscission.     

Diurnal variation in leaf extension of Salix viminalis at two nitrogen supply rates

To investigate how nitrogen supply might affect the biophysical factors controlling diurnal variation in leaf extension, pot-grown Salix viminalis L. were supplied with nitrogen at a low relative addition rate of 0.05 g N g(-1) N day(-1) (low N) or were given free access to all nutrients (high N). Leaf extension, turgor pressure, turgor after stress relaxation and the plastic extensibility of leaf tissue were determined for growing leaves every 4 h during two days of clear skies in August. Plants in the high-N treatment had a significantly higher relative growth rate, dry weight, shoot/root ratio, leaf nitrogen concentration, total leaf area, final area of single leaves and epidermal cell size than plants in the low-N treatment. The periodicity of leaf extension was similar in both treatments with high values during the afternoon and early evening, and negligible values during the night and in the early morning. The maximum rate of leaf extension was higher in high-N than in low-N plants. Leaf water potential and leaf osmotic potential decreased in the morning and increased in the afternoon with highest values during the night. Calculated values of turgor pressure showed no consistent diurnal trend and did not correlate with the rate of leaf extension. There was no consistent difference in turgor between treatments. Turgor after stress relaxation varied diurnally. The difference between turgors before and after stress relaxation also varied diurnally and was largely in phase with the diurnal pattern of leaf extension. These data are consistent with either a causal role for growth turgor (difference between turgors before and after stress relaxation) in the regulation of cell expansion, or a diurnal variation in turgors after relaxation, attributable to different capacities for cell wall loosening at different times of day. Plastic extensibility of leaf tissue showed no diurnal pattern but consistently higher values were found in high-N than in low-N plants. We conclude that the effects of nitrogen supply on leaf water relations did not limit leaf extension, but that nitrogen supply did affect processes associated with cell wall loosening and enlargement. Nitrogen supply did not affect final values of turgor after relaxation, but it presumably affected the rate at which relaxation proceeded.     

Variability in Populus leaf anatomy and morphology in relation to canopy position, biomass production, and varietal taxon

Twelve poplar (Populus) genotypes, belonging to different taxa and to the sections Aigeiros and Tacamahaca, were studied during the third growing season of the second rotation of a high density coppice culture. With the objective to highlight the relationships between leaf traits, biomass production and taxon as well as the influence of canopy position, anatomical and morphological leaf characteristics (i.e. thickness of epidermis, of palisade and spongy parenchyma layers, density and length of stomata, leaf area, specific leaf area (SLA) and nitrogen concentration) were examined for mature leaves from all genotypes and at two canopy positions (upper and lower canopy). Above ground biomass production, anatomical traits, stomatal and morphological leaf characteristics varied significantly among genotypes and between canopy positions. The spongy parenchyma layer was thicker than the palisade parenchyma layer for all genotypes and irrespective of canopy position, except for genotypes belonging to the P. deltoides × P. nigra taxon (section Aigeiros). Leaves at the upper canopy position had higher stomatal density and thicker anatomical layers than leaves at the lower canopy position. Leaf area and nitrogen concentration increased from the bottom to the top of the canopy, while SLA decreased. Positive correlations between biomass production and abaxial stomatal density, as well as between biomass production and nitrogen concentration were found. A principal component analysis (PCA) showed that genotypes belonging to the same taxon had similar anatomical characteristics, and genotypes of the same section also showed common leaf characteristics. However, Wolterson (P. nigra) differed in anatomical leaf characteristics from other genotypes belonging to the same section (section Aigeiros). Hybrids between the two sections (Aigeiros × Tacamahaca) expressed leaf characteristics intermediate between both sections, while their biomass production was low.     

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.     

Seasonal variation in the photosynthetic capacity of a willow (Salix cv. Aquatica gigantea) canopy. 1 Changes in the activity and amount of ribulose 1,5-bisphosphate carboxylase-oxygenase and the content of nitrogen and chlorophyll at different levels in the canopy

The role of ribulose 1,5-bisphosphate carboxylase-oxygenase (Rubisco) in the regulation of photosynthesis in the field was assessed by following changes in the amount and activity of Rubisco and in the amounts of chlorophyll and total nitrogen in a willow (Salix cv. Aquatica gigantea) canopy during two growing seasons. An additional aim of this study was to determine how nitrogen is partitioned in the canopy and whether Rubisco and chlorophyll serve as sources of nitrogen. During both growing seasons, the total activity and the amount of Rubisco were high in young, sun-exposed leaves and decreased as the leaves aged and became shaded. In 1985, the specific activity was low (on average 0.8 micromol mm(-1) mg(-1) Rubisco protein) compared with the values obtained in 1986 (on average 1.8 micromol min(-1) mg(-1) Rubisco protein). These differences in the specific activity of Rubisco between the two years might be associated with changes in partitioning of nitrogen. During the cool summer of 1985, about 20% of the total nitrogen was partitioned to Rubisco compared with about 13% during the warm summer of 1986. Rubisco comprised a high proportion (about 60%) of the total soluble protein throughout the 1985 growing season, whereas during the 1986 growing season, the proportion of Rubisco in the total soluble protein fraction was lower and decreased markedly as the leaves became shaded. Chlorophyll did not serve as a source of nitrogen in the canopy. On the contrary, the proportion of total nitrogen in chlorophyll increased with time, which indicates acclimation of the thylakoid membranes to decreasing irradiance. The degree of activation of Rubisco in samples collected in 1986 was high in young, sun-exposed leaves, which suggests that in these leaves, Rubisco may limit the rate of CO(2) uptake. In the shaded leaves of the lower canopy, the degree of activation of Rubisco was low, which indicates that in these leaves, photosynthesis is limited by factors other than Rubisco.     

4种柳树苗木叶的PV水分参数对水分胁迫的响应

在温室内研究了瑞典能源柳C号、2号柳、4号柳和陕北红皮柳苗木的抗旱能力。对其盆栽苗木施行水分轻度胁迫、中度胁迫、重度胁迫及正常(供水)生长4种处理,应用压力室和PV技术测定4种柳树苗木在不同处理状态下的叶组织多项水分参数,以此对其苗木的耐旱能力作综合评定。结果表明,在水分胁迫条件下,2号柳苗木叶吸收水分的能力最强;4种柳树的苗木均有一定的渗透调节和保持膨压能力,4号柳、2号柳、红柳的苗木能通过增加叶束缚水的含量来实现渗透调节,但其依靠改变苗木叶束缚水的含量来实现渗透调节的作用有限,红柳、4号柳苗木在土壤相对含水量降至61%时,2号柳苗木在土壤相对含水量降至46%时,其叶由束缚水含量所引起的渗透调节作用消失;在水分胁迫条件下,红柳、4号柳苗木通过改变叶组织细胞壁的弹性来保持其的膨压,2号柳、能源柳C号的苗木主要靠叶渗透调节来维持其的膨压。水分胁迫能增强红柳、能源柳C号、4号柳苗木保持最大膨压的能力,而降低2号柳苗木维持最大膨压的能力;4号柳能通过提高苗木叶细胞原生质忍耐脱水的能力来适应干旱,能源柳C号、红柳苗木叶细胞原生质耐脱水能力对水分胁迫的适应性差,在水分胁迫条件下其耐脱水能力减弱。通过对4种柳树苗木叶组织渗透调节和维持膨压能力的综合比较,其抗旱能力由强至弱依次为能源柳C号、2号柳、红柳和4号柳。     

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.     

Enhancement of understory productivity by asynchronous phenology with overstory competitors in a temperate deciduous forest

Some saplings and shrubs growing in the understory of temperate deciduous forests extend their periods of leaf display beyond that of the overstory, resulting in periods when understory radiation, and hence productivity, are not limited by the overstory canopy. To assess the importance of the duration of leaf display on the productivity of understory and overstory trees of deciduous forests in the north eastern United States, we applied the simulation model, BIOME-BGC with climate data for Hubbard Brook Experimental Forest, New Hampshire, USA and mean ecophysiological data for species of deciduous, temperate forests. Extension of the overstory leaf display period increased overstory leaf area index (LAI) by only 3 to 4% and productivity by only 2 to 4%. In contrast, extending the growing season of the understory relative to the overstory by one week in both spring and fall, increased understory LAI by 35% and productivity by 32%. A 2-week extension of the growing period in both spring and fall increased understory LAI by 53% and productivity by 55%.     

Calibration and assessment of seasonal changes in leaf area index of a tropical dry forest in different stages of succession

A simple measure of the amount of foliage present in a forest is leaf area index (LAI; the amount of foliage per unit ground surface area), which can be determined by optical estimation (gap fraction method) with an instrument such as the Li-Cor LAI-2000 Plant Canopy Analyzer. However, optical instruments such as the LAI-2000 cannot directly differentiate between foliage and woody components of the canopy. Studies investigating LAI and its calibration (extracting foliar LAI from optical estimates) in tropical forests are rare. We calibrated optical estimates of LAI from the LAI-2000 with leaf litter data for a tropical dry forest. We also developed a robust method for determining LAI from leaf litter data in a tropical dry forest environment. We found that, depending on the successional stage of the canopy and the season, the LAI-2000 may underestimate LAI by 17% to over 40%. In the dry season, the instrument overestimated LAI by the contribution of the woody area index. Examination of the seasonal variation in LAI for three successional stages in a tropical dry forest indicated differences in timing of leaf fall according to successional stage and functional group (i.e., lianas and trees). We conclude that when calculating LAI from optical estimates, it is necessary to account for the differences between values obtained from optical and semi-direct techniques. In addition, to calculate LAI from litter collected in traps, specific leaf area must be calculated for each species rather than from a mean value for multiple species.     

Shoot biomass growth is related to the vertical leaf nitrogen gradient in Salix canopies

Plant canopy optimization models predict that leaf nitrogen (N) distribution in the canopy will parallel the vertical light gradient, and numerous studies with many species have confirmed this prediction. Further, it is predicted that for a given canopy leaf area, a low vertical light extinction coefficient will promote rapid growth. Therefore, the ideal canopy of fast-growing plants should combine high leaf area index with a low light extinction coefficient; the latter being reflected in a flat vertical leaf N gradient throughout the canopy. Based on data from an experimental Salix stand (six varieties) grown on agricultural land in central Sweden, we tested the hypothesis that shoot growth is correlated with vertical leaf N gradient in canopies of hybrid willows bred for biomass production, which could have implications for Salix breeding. Tree improvement research requires screening of growth-related traits in large numbers of plants, but assessment of canopy leaf N gradients by chemical analysis is expensive, time-consuming and destructive. An alternative to analytical methods is to estimate leaf N gradients nondestructively with an optical chlorophyll meter (SPAD method). Here we provide a specific calibration for interpreting SPAD data measured in hybrid willows grown in biomass plantations on fertile agricultural land. Based on SPAD measurements, a significant and inverse relationship (r(2) = 0.88) was found between shoot biomass growth and vertical leaf N gradient across canopies of six Salix varieties.     

Soil temperature,gas exchange and nitrogen status of 5-year-old Norway spruce seedlings

Five-year-old Norway spruce (Picea abies (L.) Karst.) seedlings were subjected to three simulated growing seasons in controlled environment chambers. Plants were acclimated to a soil temperature of 16 degrees C during the first and third growing seasons, but were allocated at random to soil temperature treatments of 9, 13, 18 and 21 degrees C during the second growing season. Low soil temperature during the second growing season depressed stomatal conductance and photosynthetic rate (A) per unit of projected leaf area, although intercellular CO2 concentrations did not differ significantly between treatments. At all soil temperatures, total chlorophyll concentration first decreased and then increased, although the rate of increase and the final concentration increased with soil temperature, which may explain the effect of soil temperature on A. Neither chlorophyll a/b ratio nor leaf nitrogen concentration was significantly affected by soil temperature. Treatment differences disappeared during the third simulated growing season when plants were again acclimated to a soil temperature of 16 degrees C.     

Leaf photosynthetic properties in a willow (Salix viminalis and Salix dasyclados) plantation in response to fertilization

Specific leaf area (SLA), nitrogen and chlorophyll concentrations and photosynthetic characteristics were studied in upper and lower canopy leaves of Salix viminalis and S. dasyclados grown at two nutrition levels. Fertilization increased SLA and leaf mass-based nitrogen concentration in most cases. Positive effects of fertilization on leaf light-saturated photosynthetic rate (A _max ^A ) and maximum carboxylation rate (V _cmax) were not detected. Significant differences between the leaves from upper and lower canopy layers in area-based nitrogen, A _max ^A , SLA, mass-based chlorophyll, V _cmax and stomatal conductance were found for most plots. We attempted to estimate the fraction of non-photosynthetic nitrogen and found that it tended to be higher due to fertilization. Thus, the insensitivity of leaf photosynthesis to fertilization could be caused by higher proportion of non-photosynthetic nitrogen in the leaves of fertilized plots. Though leaf-level photosynthesis was not increased by fertilization, considerably higher leaf area index of fertilized plots still resulted in increased canopy carbon gain.     

Contrasting seasonal leaf habits of canopy trees between tropical dry-deciduous and evergreen forests in Thailand

We compared differences in leaf properties, leaf gas exchange and photochemical properties between drought-deciduous and evergreen trees in tropical dry forests, where soil nutrients differed but rainfall was similar. Three canopy trees (Shorea siamensis Miq., Xylia xylocarpa (Roxb.) W. Theob. and Vitex peduncularis Wall. ex Schauer) in a drought-deciduous forest and a canopy tree (Hopea ferrea Lanessan) in an evergreen forest were selected. Soil nutrient availability is lower in the evergreen forest than in the deciduous forest. Compared with the evergreen tree, the deciduous trees had shorter leaf life spans, lower leaf masses per area, higher leaf mass-based nitrogen (N) contents, higher leaf mass-based photosynthetic rates (mass-based P(n)), higher leaf N-based P(n), higher daily maximum stomatal conductance (g(s)) and wider conduits in wood xylem. Mass-based P(n) decreased from the wet to the dry season for all species. Following onset of the dry season, daily maximum g(s) and sensitivity of g(s) to leaf-to-air vapor pressure deficit remained relatively unchanged in the deciduous trees, whereas both properties decreased in the evergreen tree during the dry season. Photochemical capacity and non-photochemical quenching (NPQ) of photosystem II (PSII) also remained relatively unchanged in the deciduous trees even after the onset of the dry season. In contrast, photochemical capacity decreased and NPQ increased in the evergreen tree during the dry season, indicating that the leaves coped with prolonged drought by down-regulating PSII. Thus, the drought-avoidant deciduous species were characterized by high N allocation for leaf carbon assimilation, high water use and photoinhibition avoidance, whereas the drought-tolerant evergreen was characterized by low N allocation for leaf carbon assimilation, conservative water use and photoinhibition tolerance.     

Fall frost resistance in willows used for biomass production. II. Predictive relationships with sugar concentration and dry matter content

The accumulation of sugars and dry matter in stems in fall was examined in relation to frost hardening in eight willow clones (six clones of Salix viminalis L. and one clone each of S. viminalis x S. schwerenii E. Wolf and S. dasyclados Wimm.). Evidence is presented that three sources of variation in fall frost resistance among the eight clones could be assessed from an analysis of stem composition. First, the pre-hardening value of frost resistance could be assessed from the total sugar concentration. Second, the start of induction of apical growth cessation and hence frost hardening could be distinguished by a stepwise increase in sucrose-to-glucose ratio. Third, the progress of frost hardening during its first phase could be followed from a proportional rise in total sugar concentration and, even more accurately, from a proportional rise in dry-to-fresh weight ratio. In contrast, the second phase of frost hardening was largely uncoupled from sugar and dry matter accumulation. Raffinose and sucrose accumulation seemed to be under differential environmental controls. Sucrose accumulation started with the initiation of growth cessation controlled by photoperiod, whereas raffinose accumulation started with falling temperatures later on. Starch reserves that built up in stems in early fall were partially mobilized later on to support sugar accumulation. In contrast to stems, leaves did not exhibit a preferential accumulation of sucrose in fall.     

Transpiration and forest structure in relation to soil waterlogging in a Hawaiian montane cloud forest

Transpiration, leaf characteristics and forest structure in Metrosideros polymorpha Gaud. stands growing in East Maui, Hawaii were investigated to assess physiological limitations associated with flooding as a mechanism of reduced canopy leaf area in waterlogged sites. Whole-tree sap flow, stomatal conductance, microclimate, soil oxidation-reduction potential, stand basal area and leaf area index (LAI) were measured on moderately sloped, drained sites with closed canopies (90%) and on level, waterlogged sites with open canopies (50-60%). The LAI was measured with a new technique based on enlarged photographs of individual tree crowns and allometric relationships. Sap flow was scaled to the stand level by multiplying basal area-normalized sap flow by stand basal area. Level sites had lower soil redox potentials, lower mean stand basal area, lower LAI, and a higher degree of soil avoidance by roots than sloped sites. Foliar nutrients and leaf mass per area (LMA) in M. polymorpha were similar between level and sloped sites. Stomatal conductance was similar for M. polymorpha saplings on both sites, but decreased with increasing tree height (r(2) = 0.72; P < 0.001). Stand transpiration estimates ranged from 79 to 89% of potential evapotranspiration (PET) for sloped sites and from 28 to 51% of PET for level sites. Stand transpiration estimates were strongly correlated with LAI (r(2) = 0.96; P < 0.001). Whole-tree transpiration was lower at level sites with waterlogged soils, but was similar or higher for trees on level sites when normalized by leaf area. Trees on level sites had a smaller leaf area per stem diameter than trees on sloped sites, suggesting that soil oxygen deficiency may reduce leaf area. However, transpiration per unit leaf area did not vary substantially, so leaf-level physiological behavior was conserved, regardless of differences in tree leaf area.     

基于土壤水分承载力的林分密度计算与调控——以六盘山华北落叶松人工林为例

立地水分条件决定的植被承载力是干旱缺水地区森林合理经营的重要依据。考虑到干旱缺水地区的森林蒸散耗水在水分输出中占据绝对主导地位,其大小直接与叶面积指数(LAI)相关,将林冠LAI在生长季一段时间内的最大值(LAImax)作为植被承载力(LAIc)的量化指标,利用冠层分析仪(LAI-2000),在六盘山香水河小流域和叠叠沟小流域的44个华北落叶松人工林样地,实测了冠层LAI的季节动态变化,研究了生长季内LAImax与林分断面积、郁闭度、平均树高、密度等常用林分结构指标的关系。结果表明:LAImax与林分不同结构指标均呈幂函数关系,其决定系数(R2)依次为0.84、0.82、0.56、0.47,说明能同时反映林分密度和树体大小的林分断面积与林冠LAI相关最紧密。将LAImax与林分断面积的幂函数关系嵌入了林分平均胸径与林分密度和林龄关系的模型,用以描述LAImax与林龄和密度的关系,并利用样地实测数据拟合了模型参数。拟合建立的模型对所有样地的LAImax的计算值与实测值的相对误差平均为8.6%(0%20.4%),能较好地描述LAI与林龄和密度的关系。利用此模型,进一步导出了能依据给定的LAIc,简捷计算出不同林龄时的可承载林分密度的模型,从而为基于立地水分植被承载力的林分密度管理和森林多功能经营等提供技术支持。     

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.