华山松、油松不同叶龄针叶光合特性对林窗式疏伐的响应

[目的]通过对比研究华山松和油松不同叶龄针叶形态、光合速率、叶绿素和叶氮含量对林窗式疏伐的响应,探讨林窗大小、叶龄和树种对林窗式疏伐后叶光合特性响应的影响。[方法]2008年在30年生华山松和油松混交人工林中,设置对照、小林窗(80 m2)和中林窗(110 m2)处理,2013年以对照样地内和林窗边缘华山松和油松为研究对象,测定其当年生、1年生和2年生叶比叶重(LMA)、单位叶面积叶绿素a(Chl a)、叶绿素b含量(Chl b)、氮含量(N)、最大净光合速率(Amax)、光合氮利用效率(PNUE)等指标。[结果]小林窗和中林窗疏伐对叶光合特性影响一致,均对当年生叶影响不显著,但显著改变了1年生和2年生叶绿素含量、N和PNUE,提高了其Amax。华山松叶LMA、叶绿素含量、N、Amax均低于油松,两树种多年生叶对林窗式疏伐的响应也明显不同:林窗边缘华山松1年生和2年生叶LMA和N与对照差异不显著,但叶绿素含量低于对照,PNUE和Amax显著高于对照;林窗边缘油松1年生和2年生叶LMA和PNUE与对照差异不显著,但N和Amax显著高于对照。[结论]80 110 m2林窗疏伐显著改变叶光合特性,提高其光合潜力;林窗疏伐影响1年生和2年生叶光合特性,对当年生叶影响不显著;华山松和油松对林窗式疏伐的响应特征存在差异。     

Functional relationships between leaf structure and photosynthetic traits as modulated by irradiance and nutrient availability in a sclerophyllous and a non-sclerophyllous mediterranean oak species

Plasticity of structural and physiological leaf traits elicited by irradiance and soil nutrients was investigated in two sympatric mediterranean oaks: a sclerophyllous (Quercus suber L.) and a non-sclerophyllous species (Q. canariensis Willd.). Seedlings were grown for 2 years in pots in a 2-way crossed factors design. Leaf mass-to-area ratio (LMA) and nitrogen were recorded, and photosynthetic capacity (i.e. the apparent maximal carboxylation rate by rubisco, V _cmax) was derived from response curves of net CO₂ assimilation (A) versus intercellular CO₂ mol fraction (C _i). Structural equation modelling was applied to the data for disentangling the complex correlation structure between variables. The two species differed significantly in photosynthetic nitrogen use efficiency (PNUE). They displayed the expected responses to irradiance, with large increases in LMA, V _cmax and nitrogen per unit leaf area and decreases in mass-based nitrogen content. Nutrient availability modulated severely leaf N content (mass- and area-based) and mass-based maximal carboxylation rate, but not the plastic response of all these parameters to irradiance. Irradiance primarily modulated leaf structure (LMA), and secondarily nitrogen content, while nutrient availability modulated directly nitrogen content. Nitrogen content in turn had a severe impact on mass-based photosynthetic capacity. It is concluded that in young trees solely leaf structure displayed irradiance-elicited plasticity. This plasticity was not modulated by nutrient availability and was similar in a sclerophyllous and a non-sclerophyllous species.     

Leaf-level responses to light in two co-occurring <Emphasis Type="Italic">Quercus</Emphasis> (<Emphasis Type="Italic">Quercus ilex</Emphasis> and <Emphasis Type="Italic">Quercus suber</Emphasis>): leaf structure,chemical composition and photosynthesis

We studied morphological, biochemical and physiological leaf acclimation to incident Photon-Photosynthetic-Flux-Density (PPFD) in Quercus ilex (holm oak) and Quercus suber (cork oak) at Mediterranean evergreen oak woodlands of southern Portugal. Specific leaf area (SLA) decreased exponentially with increasing PPFD in both species. Q. ilex had lower SLA values than Q. suber. Leaf nitrogen, cellulose and lignin concentration (leaf area-based) scaled positively with PPFD. Maximum rate of carboxylation (V_cmax), capacity for maximum photosynthetic electron transport (J_max), rate of triose-P utilization (V_TPU) and the rate of nonphotorespiratory light respiration (Rd) were also positively correlated with PPFD in both Quercus species, when expressed in leaf area but not on leaf mass basis. Q suber showed to have higher photosynthetic potential (V_cmax, J_max^m and V_TPU^m) and a higher nitrogen efficient nitrogen use than Q.ilex. Leaf chlorophyll concentration increased with decreasing PPFD, improving apparent quantum use efficiency (Φ) in both Quercus species. We concluded that, in Q.ilex and Q.suber, leaf structural plasticity is a stronger determinant for leaf acclimation to PPFD than biochemical and physiological plasticity.     

Leaf photosynthetic responses and related nitrogen changes associated with crown reclosure after thinning in a young Chamaecyparis obtusa stand

In order to quantify the effects of thinning on biochemical photosynthesis parameters and changes in leaf nitrogen contents associated with the process of crown reclosure, the maximum rate of carboxylation (V _cmax), the leaf nitrogen concentration per unit area (N _a), and the photosynthetic photon flux density (PPFD) were measured at four crown heights in both thinned (1500 trees ha⁻¹) and unthinned control (3000 trees ha⁻¹) stands of ten-year-old Chamaecyparis obtusa (36°3′N, 140°7′E) trees during four consecutive growing seasons after thinning. Thinning increased V _cmax in the lower and middle crowns in the first year after thinning, and leaves in the lower crown of the thinned stand maintained high V _cmax for four years, whereas they abscised in the second year in the control stand. Significant increases in V _cmax were detected even in the upper crowns of trees in the thinned stand in the second year. Thinning did not affect N _a at any of the crown positions in the first year, but significantly increased N _a in the middle crowns from the second year after thinning. Thus, the redistribution of nitrogen between leaves, driven by increases in light and nutrient availability due to the 50% thinning, appears to have enhanced photosynthetic rates in the thinned stand. Thinning also significantly affected the slope of the linear relationship between N _a and V _cmax initially after thinning, but its effect on this relationship was negligible after the second year. These quantitative results may be used to simplify the estimation of the likely effects of management practices on carbon fixation in forest canopies.     

Photosynthetic characteristics of Fagus sylvatica and Quercus robur established for stand conversion from Picea abies

Efforts in Europe to convert Norway spruce (Picea abies) plantations to broadleaf or mixed broadleaf-conifer forests could be bolstered by an increased understanding of how artificial regeneration acclimates and functions under a range of Norway spruce stand conditions. We studied foliage characteristics and leaf-level photosynthesis on 7-year-old European beech (Fagus sylvatica) and pedunculate oak (Quercus robur) regeneration established in open patches and shelterwoods of a partially harvested Norway spruce plantation in southwestern Sweden. Both species exhibited morphological plasticity at the leaf level by developing leaf blades in patches with an average mass per unit area (LMA) 54% greater than of those in shelterwoods, and at the plant level by maintaining a leaf area ratio (LAR) in shelterwoods that was 78% greater than in patches. However, we observed interspecific differences in photosynthetic capacity relative to spruce canopy openness. Photosynthetic capacity (A₁₆₀₀, net photosynthesis at a photosynthetic photon flux density of 1600 μmol photons m⁻² s⁻¹) of beech in respect to the canopy gradient was best related to leaf mass, and declined substantially with increasing canopy openness primarily because leaf nitrogen (N) in this species decreased about 0.9 mg g⁻¹ with each 10% rise in canopy openness. In contrast, A₁₆₀₀ of oak showed a weak response to mass-based N, and furthermore the percentage of N remained constant in oak leaf tissues across the canopy gradient. Therefore, oak photosynthetic capacity along the canopy gradient was best related to leaf area, and increased as the spruce canopy thinned primarily because LMA rose 8.6 g m⁻² for each 10% increase in canopy openness. These findings support the premise that spruce stand structure regulates photosynthetic capacity of beech through processes that determine N status of this species; leaf N (mass basis) was greatest under relatively closed spruce canopies where leaves apparently acclimate by enhancing light harvesting mechanisms. Spruce stand structure regulates photosynthetic capacity of oak through processes that control LMA; LMA was greatest under open spruce canopies of high light availability where leaves apparently acclimate by enhancing CO₂ fixation mechanisms.     

An integrated model for predicting maximum net photosynthetic rate of cocksfoot (Dactylis glomerata) leaves in silvopastoral systems

Net light-saturated photosynthetic rate (A_max) of field grown cocksfoot (Dactylis glomerata L.) leaves in a radiata pine (Pinus radiata D. Don) silvopastoral system (Canterbury, New Zealand) was measured at different times under severe shade (85–95 μmol m^–2 s^–1 photosynthetic photon flux density, PPFD) and in full sunlight (1900 μmol m^–2 s^–1 PPFD). The aim was to integrate individual functions for A_max against air temperature (2 to 37 oC), water status, expressed as pre-dawn leaf water potential (ψ_lp) (-0.01 to −1.6 MPa), herbage nitrogen (N) (1.5 to 5.9%), regrowth duration (20 to 60 days) and time under shade (1 to 180 min) into a multiplicative model. The highest A_max value obtained was 27.4 μmol CO₂ m^–2 s^–1 in non-limiting conditions with full sunlight. This value was defined as standardised dimensionless A_maxs = 1 for comparison of factor effects. The canopy temperature of the cocksfoot sward was up to 7.4 oC cooler than air temperature for plants under shade. Therefore, canopy temperature was used to predict A_max. The only interaction was between time under severe shade (5% of the open PPFD) and water stress (ψ_lp = −0.4 to −1.3 MPa) and this was included in the model. Validation of this model indicated 78% of the variation in A_max could be accounted for using these five factors by the addition of the interaction function. This model could be used to assist the prediction of pasture growth in silvopastoral systems through incorporation into a canopy photosynthesis model. This revised version was published online in June 2006 with corrections to the Cover Date.     

Photosynthetic response to green crown pruning in young plantation-grown Eucalyptus pilularis and E. cloeziana

The loss of foliage through pruning of live branches may reduce tree growth or it may be compensated by photosynthetic up-regulation of the remaining crown. Here, the changes in light-saturated photosynthesis following pruning to remove 50% of green crown length were examined in 4-year-old Eucalyptus pilularis Sm. and Eucalyptus cloeziana F. Muell. trees. The objectives of the study were to: (1) compare leaf-level physiological (light-saturated photosynthesis (A_max), stomatal conductance (g), transpiration (T), dark respiration (R_d), quantum yield (Φ), light compensation point (Γ), water-use efficiency (WUE), nitrogen-use efficiency (NUE)) traits in species with contrasting crown dynamics and structure, (2) examine the effect of crown position on these traits, and (3) examine the effect of pruning on A_max, g, T, WUE, NUE, leaf N and P concentrations and specific leaf area (SLA). Prior to pruning there were no differences in R_d, Γ and Φ between E. pilularis and E. cloeziana but differences in A_max, T, g, leaf N, leaf P, WUE, NUE and SLA. Whereas the rate of physiological processes (A_max, T, and g) and leaf N and P concentrations increased with crown height, R_d, Γ, Φ and SLA declined along this vertical gradient, except in the upper crown of E. cloeziana where A_max, T and g were not different to the lower crown. No up-regulation of photosynthesis or changes in leaf physiology occurred between 6 and 13 months after pruning in either species. The results provide an important basis for modelling pruning effects in process-based tree growth models.     

Nitrogen fertilization of black walnut (Juglans nigra L.) during plantation establishment. Physiology of production

Physiological mechanisms by which nitrogen (N) fertilization affects growth and development in temperate deciduous forest trees are not clearly understood, especially under intensive silvicultural systems. Grafted, Tippecanoe 1 cultivar black walnut (Juglans nigra L.) trees were grown in an intensively managed plantation in west-central Spain and subjected to six, fixed-nutrient-ratio complete fertilizer treatments (defined as 0, 25, 50, 75, 150, and 300 g N tree^?1) delivered via daily fertigation. Leaf chemistry and morphology were evaluated from June to September, and gas exchange was measured in July. Specific leaf mass, leaflet nitrogen (N), and chlorophyll concentrations varied over the course of the growing season, yet consistently increased with increasing fertilization. Net photosynthesis at ambient (A _net) and light-saturated (A _max) conditions increased from the unfertilized control to lowest treatment (25 g N) but did not increase at higher fertilizer rates. Photosynthetic N and chlorophyll use efficiencies decreased with increasing fertilization, but photosynthetic phosphorus and water use efficiencies increased. Transpiration rates and dark respiration were not significantly affected by treatment. Overall, the lowest fertilizer treatment (25 g N) had the greatest photosynthetic efficiency. Interactions between N and other nutrients with increasing fertilizer application suggested potential for nutrient imbalances at high fertilization rates. Our results provide a physiological justification for the use of low-to-moderate fertilization as an efficient strategy to promote black walnut plantation establishment under intensive cultural systems.     

Morphological and physiological adjustment to N and P fertilization in nutrient-limited Metrosideros polymorpha canopy trees in Hawaii

Leaf-level studies of Metrosideros polymorpha Gaud. (Myrtaceae) canopy trees at both ends of a substrate age gradient in the Hawaiian Islands pointed to differential patterns of adjustment to both nutrient limitation and removal of this limitation by long-term (8-14 years) nitrogen (N), phosphorus (P) and N + P fertilizations. The two study sites were located at the same elevation, had similar annual precipitation, and supported forests dominated by M. polymorpha, but differed in the age of the underlying volcanic substrate, and in soil nutrient availability, with relatively low N at the young site (300 years, Thurston, Hawaii) and relatively low P at the oldest site (4,100,000 years, Kokee, Kauai). Within each site, responses to N and P fertilization were similar, regardless of the difference in soil N and P availability between sites. At the young substrate site, nutrient addition led to a larger mean leaf size (about 7.4 versus 4.8 cm2), resulting in a larger canopy leaf surface area. Differences in foliar N and P content, chlorophyll concentrations and carboxylation capacity between the fertilized and control plots were small. At the old substrate site, nutrient addition led to an increase in photosynthetic rate per unit leaf surface area from 4.5 to 7.6 micromol m(-2) s(-1), without a concomitant change in leaf size. At this site, leaves had substantially greater nutrient concentrations, chlorophyll content and carboxylation capacity in the fertilized plots than in the control plots. These contrasting acclimation responses to fertilization at the young and old sites led to significant increases in total carbon gain of M. polymorpha canopy trees at both sites. At the young substrate site, acclimation to fertilization was morphological, resulting in larger leaves, whereas at the old substrate site, physiological acclimation resulted in higher leaf carboxylation capacity and chlorophyll content.     

Validation of a canopy photosynthesis model for cocksfoot pastures grown under different light regimes

Daily net canopy photosynthesis (P _n) was predicted for cocksfoot (Dactylis glomerata L.) canopies grown under different light regimes by integration of a leaf photosynthesis model developed for the light-saturated photosynthetic rate (P _max), photosynthetic efficiency (α) and the degree of curvature (θ) of the leaf light–response curve. When shade was the only limiting factor, the maximum P _n (P _nmax) was predicted to decrease approximately linearly from 33.4 g CO₂ m⁻² d⁻¹ to zero as photosynthetic photon flux density (PPFD) fell from full sunlight (1800 μmol m⁻² s⁻¹ PPFD) to 10% of this in a fluctuating light regime. It was also predicted that at 50% transmissivity P _nmax was higher for a continuous light regime (10.4 g CO₂ m⁻² d⁻¹) than for a fluctuating light regime with the same intensity (8.4 g CO₂ m⁻² d⁻¹). The canopy photosynthesis model was then used to predict dry matter (DM) production for cocksfoot field grown pastures under a diverse range of temperature, herbage nitrogen content and water status conditions in fluctuating light regimes. This prediction required inclusion of leaf area index and leaf canopy angle from field measurements. The model explained about 85% of the variation in observed cocksfoot DM production for a range from 6 to 118 kg DM ha⁻¹ d⁻¹. The proposed model improves understanding of pasture growth prediction through integration of relationships between shade limitations in fluctuating light regimes and other environmental factors that affect the canopy photosynthetic rate of cocksfoot pastures in silvopastoral systems.     

Seasonal changes in leaf nitrogen pools in two Salix species

Leaf nitrogen distribution pattern was studied four times during the growing season in a 2-year-old Salix viminalis L. and Salix dasyclados Wimm. plantation in Estonia. We measured the vertical distributions of leaf nitrogen concentration, dry mass, leaf area and light environment (as fractional transmission of diffuse irradiance, a(d)) in the canopy. The light-independent nitrogen pool was evaluated as the intercept of the leaf nitrogen concentration versus a(d) relationship, and the nondegradable nitrogen pool was evaluated as the nitrogen remaining in abscised leaves. A strong vertical gradient of mass-based leaf nitrogen concentration was detected at the beginning of the growing season, and decreased steadily during canopy development. This decline had at least three causes: (1) the amount of nitrogen in the foliage was larger at the beginning of the growing season than at the end of the growing season, probably because of pre-existing root systems; (2) with increasing leaf area index (LAI) during the growing season, the proportion of leaf nitrogen in total canopy nitrogen that could be redistributed (light-dependent nitrogen pool) decreased; and (3) the photosynthetic photon flux density gradient inside the canopy changed during the season, most probably because of changes in leaf area and leaf angle distributions. Total canopy nitrogen increased almost proportionally to LAI, whereas the light-dependent nitrogen pool had a maximum in August. Also, the proportion of the light-dependent nitrogen pool in the total canopy nitrogen decreased steadily from 65.2% in June to 17.2% in September in S. dasyclados and from 63.3 to 15.1% in S. viminalis. The degradable nitrogen pool was always bigger than the light-dependent nitrogen pool.     

Low light acclimation in five temperate broad-leaved tree species of different successional status: the significance of a shade canopy

?Context

Tree species differ largely in their capability to produce characteristic shade leaves with effective morphological and physiological acclimation to low light.

?Aims

By examining the sun/shade leaf differentiation in leaf morphology, foliar nitrogen and photosynthetic capacity in five temperate tree species of different successional status, we aimed at identifying those leaf traits that determine the development of a typical shade crown with low light-acclimated leaves.

?Methods

Leaf morphology, foliar N content, photosynthetic capacity (V _cmax, J _max and A _max) and leaf dark respiration (R _d) were measured in the canopies of 26 adult trees of Fraxinus, Acer, Carpinus, Tilia and Fagus species.

?Results

Six traits (the sun/shade leaf differentiation in specific leaf area, leaf size, A _max per leaf area or per mass, photosynthetic N use efficiency and R _d) were found to characterise best the degree of low light acclimation in shade leaves. All five species exhibited certain modifications in leaf morphology and/or physiology in response to low light; Fagus sylvatica showed the highest and Fraxinus excelsior the lowest shade leaf acclimation.

?Conclusions

Our results indicate that the five early/mid- to late-successional species have developed species-specific low light acclimation strategies in their shade crowns which differ in terms of the relative importance of leaf morphological and physiological acclimation.     

Does strategy of resource acquisition in tropical woody species vary with life form,leaf texture,and canopy gradient?

In order to test whether the strategy of resource acquisition varies with life form, leaf texture and canopy gradient, we measured light-saturated net photosynthetic capacity (A _max-mass) and leaf nitrogen and phosphorus concentrations (N _mass and P _mass) for 127 woody species of understory (small shrubs and tree seedlings) and 47 woody species of canopy (large shrubs and tree adults) in a tropical montane rain forest of Hainan Island, South China. Photosynthetic nitrogen use efficiency (PNUE) and photosynthetic phosphorus use efficiency (PPUE) were studied by taking into account functional groups, which classified by either life form (FG1) or leaf texture (FG2). For FG1, there were significant (P < 0.05) differences between trees and shrubs in A _mass, N _mass and P _mass, but not in PNUE and PPUE; whereas for FG2, there were significant (P < 0.05) differences between the papery-leaved species and those with leathery leaves in the measured leaf traits inclusive of PNUE and PPUE. Within the same classification systems, significant allometric scaling relationships were found in the leaf trait pairs of A _mass-N _mass and A _mass-P _mass. Considered separately, the slope and y-intercept of the linear regression between A _mass and N _mass did not differ among functional groups nor between understory and canopy, but those for the linear regression between A _mass and P _mass differed significantly (P = 0.001) between understory and canopy species regardless of functional groups. The results of phylogenetic comparative analyses were in accordance with the observed positive scaling relationships. The overall results indicate that there are no fundamentally different nitrogen capture strategies in tropical woody species regardless of their life form, leaf texture and canopy gradient. However, the strategies of phosphorus acquisition of tropical woody species differ with canopy layer increases. This variation may correspond to the special soil conditions in this ecosystem, as phosphorus is an element limiting plant growth in tropical areas.     

Responses to temperature and shade in Abies alba seedlings from diverse provenances

Abstract

Intraspecific variability in responses to temperature and shade was studied at Champenoux, north-eastern France, with seedlings from five Polish provenances of silver fir (Abies alba Mill.). Acclimation of photosynthesis to temperature was investigated in seedlings exposed to 10, 25 and 35°C in a climate chamber for 1 week. During two growth seasons, a population of seedlings was grown in the nursery under four different irradiance regimens: 100, 48, 18 and 8% of natural irradiance. Maximum carboxylation rate (V _cmax), maximum light driven electron flow (J _max) and maximum net carbon dioxide assimilation rate (A _max) measured at 25°C increased with population altitude. One week of exposure to 35°C caused discoloration and massive needle shedding. After 2 years’ acclimation to different levels of irradiance, a significant interprovenance variability was evidenced in growth, total biomass, biomass allocation and photosynthetic performance. This study provided evidence for the existence of functional variation among the examined provenances.     

Effects of long-term CO2 and temperature elevation on crown nitrogen distribution and daily photosynthetic performance of Scots pine

Single Scots pines (Pinus sylvestris L.), aged 20–25 years, were grown in open-top chambers and exposed to elevated temperature (Elev. T), elevated CO₂ (Elev. C) and a combination of elevated CO₂ and temperature (Elev. C + T) for 4 years. The vertical distribution of needle nitrogen concentration was measured simultaneously with gas exchange of attached shoots. Based on the measurements, the dependencies on needle nitrogen concentrations of four photosynthetic parameters, i.e., RuP₂ (ribulose 1,5-bisphosphate)-saturated rate of carboxylation (V_cmax), maximum potential electron transport (J_max), the rate of respiration in the light (R_d) and light-use-efficiency factor (δ), were determined. Using a crown multilayer model, the performance of daily crown photosynthesis in Scots pine was predicted. Compared to the control treatment, the mean concentration of nitrogen in the foliage decreased by 20% and by 17% for trees grown under Elev. C and under Elev. C + T, respectively, but increased by 4% for trees grown under Elev. T. However, the total content of foliage nitrogen per unit ground area increased by 25% for trees grown under Elev. C, by 19% for trees grown under Elev. C + T and by 6% for trees grown under Elev. T; these were due to the increase in the total needle area index. Regressions showed that the foliage grown under Elev. C and Elev. C + T had steeper slopes representing the responses of V_cmax, R_d and δ to leaf nitrogen concentrations, while Elev. C + T and Elev. T had steeper slopes representing the response of J_max to needle nitrogen concentrations. Predictions showed that, on a typical sunny day, the daily total of crown photosynthesis increased 22% and 27%, separately for Elev. C and Elev. C + T, and by only 9% for Elev. T alone. Furthermore, the increased daily crown photosynthesis, resulting from treatments involving elevated CO₂, can be attributed mainly to an increase in the ambient CO₂ concentration and the needle area index, while modification of the intrinsic photosynthetic capacity had only a marginal effect. Based on the current pattern of crown nitrogen allocation, the prediction showed also that the relationship between daily crown photosynthesis and crown nitrogen content was strongly dependent on the daily incident PAR and air temperature. The CO₂-elevated treatments led to an increase in the sensitivity of daily crown photosynthesis to changes in crown nitrogen content, daily incident PAR and temperature, while the temperature-elevated treatment had the opposite effect on the sensitivity.     

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.     

Temporal and spatial patterns of net photosynthesis in 12-year-old loblolly pine five growing seasons after thinning

Physiological parameters were measured under natural light conditions and needle orientation from towers and walkways erected in the canopy of a loblolly pine (Pinus taeda L.) plantation. Four silvicultural treatments were randomly assigned to the twelve plots in the fall of 1988. Plots were thinned to a density of 731 trees per hectare or left unthinned, at a density of 2990 trees per hectare. The plots were left unfertilized or fertilized with 744 kg/ha of diammonium triple superphosphate was applied. During the fifth growing season (1993) following thinning and fertilization, needle level physiology was not different with respect to the thinning treatment for fertilized or unfertilized plots. In contrast, upper crown levels within the fertilized and unfertilized plots had significantly higher light levels and photosynthetic rates than lower crown foliage. Light levels were greater in the thinned, fertilized plots than in the unthinned, fertilized plots. In contrast, no effect of thinning on canopy light levels was found in the unfertilized plots. Within crown variation in photosynthesis was strongly dependent on canopy light levels. A strong interaction of canopy level with thinning was apparent for net photosynthesis. Loblolly pine, being a shade intolerant species, showed only small physiological differences between needles from different parts of the crown. Because of the variability found in this study, more extensive sampling is needed to correctly describe the physiology of a forest canopy with adequate precision.     

Within-canopy nitrogen and photosynthetic gradients are unaffected by soil fertility in field-grown Eucalyptus globulus

A significant and well-supported hypothesis is that increased growth following nitrogen (N) fertilization is a function of the relationships among photosynthesis, tissue N content and the light environment-specifically, the within-canopy allocation of N among leaves and the within-leaf allocation of N between Rubisco and chlorophyll. We tested this hypothesis in a field trial that included annual applications of N,P,K fertilizer (from planting) to a Eucalyptus globulus Labill. plantation growing on uniform leached sands. Growth of 4-year-old E. globulus increased significantly in response to fertilization. Leaf N and phosphorus concentrations were 0.1-0.5 g m(-2) and 0.4-0.5 g m(-2) higher in fertilized trees compared to unfertilized trees, respectively. Stomatal conductance (g(s)) at the maximum photosynthetic rate (A(max)) was between 0.2 and 0.4 mol m(-2) s(-1) higher in fertilized trees, but A(max) and the concentration of Rubisco (Rub(a)) were unaffected by fertilization. This seeming paradox, where there was no response of A(max) to fertilization despite increases in g(s) and leaf N concentration, was explained by reduced in vivo specific activity of Rubisco in fertilized trees. Acclimation to light, measured by redistribution of N between Rubisco and chlorophyll, was unaffected by fertilization. Distribution of leaf N followed irradiance gradients, but A(max) did not. Maximum photosynthetic rate was correlated with leaf N concentration only in unfertilized trees. These findings indicate that the relationships among photosynthesis, N and the light environment in E. globulus are affected by N,P,K fertilization.     

Response of Pinus koraiensis seedling growth to different light conditions based on the assessment of photosynthesis in current and one-year-old needles

As one of the three major five-leaved pines in the northern hemisphere, Pinus koraiensis is the most important dominant tree species in the natural mixed-broadleaved Korean pine forests. However, the regeneration of P koraiensis under the canopy of secondary forest stands is poor because of the light limitation. This study was conducted to understand how P koraiensis seedlings adapt to different light intensities and what would be the optimum light level for their establishment and growth. Three repetition plots with four light intensities （15%, 30%, 60% and 100% of the natural incident irradiances, achieved by suspending layers of black nylon net above and surrounding the plots） were set up under natural climate conditions in a montane region in eastern Liaoning Province, Northeast China. A total of 80 P koraiensis seedlings with similar height and root collar diameter were transplanted into four plots. After one year of acclimation to the specific light conditions, the seasonal variations of the photosynthetic variables and needle traits of the current and one-year-old needles, and the growth parameters were observed under four light intensities. The results indicated that： （1） The seedling at 60% treatment exhibited the greatest growth, which agreed with the response of the light-saturated photosynthetic rates （Amax） and the dark respiration rate （Rd） in the current and one-year-old needles, i.e., Rd at 60% treatment was significantly lower than that at 100% treatment, but Amax did not differ between the seedlings at 100% and 60% treatments. （2） The P. koraiensis seedlings have a certain photosynthetic plasticity to adapt the light conditions by adjusting their needle traits and regulating the physiological processes, because Amax, Rd, light saturation point and compensation point, the needle mass area, needle nitrogen and chlorophyll contents were significantly （p〈0.05） correlated with the light intensities. Especially, Am,x at 100% and 60% treatments was significantly higher （p〈0.05） than that at 30% and 15% treatments for both current and one-year-old needles. （3） The needles of different ages played a commutative role during the growing season, i.e., the one-year-old needles played a major role for the photosynthesis in the early growing season; the current year needles did in the later growing season. This ensured the effective photosynthesis throughout the growing season. These findings suggest that P. koraiensis is the in-between heliophilous and shade-tolerant tree species at least for the seedlings up to 8 years.     

Changes in leaf water use after removal of leaf lower surface hairs on <Emphasis Type="Italic">Mallotus macrostachyus</Emphasis> (Euphorbiaceae) in a tropical secondary forest in Malaysia

Leaf hairs may assist in maintaining high leaf water use efficiency in tropical secondary forest tree species. We compared leaf temperature, transpiration, photosynthesis and water use efficiency between hairy and depilated leaves in Mallotus macrostachyus (Euphorbiaceae), to determine the role of leaf hair in leaf water use efficiency (WUE) in tropical degraded secondary forest in Malaysia. Measurements were made on five mature individuals growing in sun-exposed conditions and five in shaded conditions. The hair dry weight per unit leaf area was significantly greater in sun leaves than in shade leaves. The transpiration rate (Tr_max) of depilated leaves in sun-exposed conditions was slightly higher than in hairy leaves in both morning and afternoon measurements. In contrast, Tr_max in the shade leaves was almost identical in hairy and depilated leaves. Leaf stomatal conductance (g _s) in the morning showed almost the same value among leaf types and light conditions. In the afternoon, g _s slightly decreased from the morning values in both sun and shade conditions. In the morning, the leaf water use efficiency (A _max/Tr_max) in both conditions did not differ significantly between hairy and depilated leaves. However, in the afternoon, WUE in the depilated leaves was significantly lower than in hairy leaves in sun-exposed conditions. These observations suggest that leaf hairs in M. macrostachyus contribute to the high leaf water use efficiency in drought conditions, such as high vapor pressure deficit experienced at midday in degraded tropical secondary forests.