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.     

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.     

Leaf photosynthetic traits of 14 tropical rain forest species in relation to leaf nitrogen concentration and shade tolerance

Variability of leaf traits related to photosynthesis was assessed in seedlings from 14 tree species growing in the tropical rain forest of French Guiana. Leaf photosynthetic capacity (maximum rate of carboxylation and maximum rate of electron transport) was estimated by fitting a biochemical model of photosynthesis to response curves of net CO2 assimilation rate versus intercellular CO2 mole fraction. Leaf morphology described by leaf mass per unit leaf area (LMA), density and thickness, as well as area- and mass-based nitrogen (N) and carbon (C) concentrations, were recorded on the same leaves. Large interspecific variability was detected in photosynthetic capacity as well as in leaf structure and leaf N and C concentrations. No correlation was found between leaf thickness and density. The correlations between area- and mass-based leaf N concentration and photosynthetic capacity were poor. Conversely, the species differed greatly in relative N allocation to carboxylation and bioenergetics. Principal component analysis (PCA) revealed that, of the recorded traits, only the computed fraction of total leaf N invested in photosynthesis was tightly correlated to photosynthetic capacity. We also used PCA to test to what extent species with similar shade tolerances displayed converging leaf traits related to photosynthesis. No clear-cut ranking could be detected among the shade-tolerant groups, as confirmed by a one-way ANOVA. We conclude that the large interspecific diversity in photosynthetic capacity was mostly explained by differences in the relative allocation of N to photosynthesis and not by leaf N concentration, and that leaf traits related to photosynthetic capacity did not discriminate shade-tolerance ranking of these tropical tree species.     

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.     

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.     

Photosynthetic light acclimation in peach leaves: importance of changes in mass:area ratio, nitrogen concentration, and leaf nitrogen partitioning

Photosynthetic light acclimation of leaves can result from (i) changes in mass-based leaf nitrogen concentration, Nm, (ii) changes in leaf mass:area ratio, Ma, and (iii) partitioning of total leaf nitrogen among different pools of the photosynthetic machinery. We studied variations in Nm and Ma within the crowns of two peach (Prunus persica L. Batsch) trees grown in an orchard in Portugal, and one peach tree grown in an orchard in France. Each crown was digitized and a 3-D radiation transfer model was used to quantify the intra-crown variations in time-integrated leaf irradiance, . Nitrogen concentration, leaf mass:area ratio, chlorophyll concentration, and photosynthetic capacity were also measured on leaves sampled on five additional peach trees in the orchard in Portugal. The data were used to compute the coefficients of leaf nitrogen partitioning among carboxylation, bioenergetics, and light harvesting pools. Leaf mass:area ratio and area-based leaf nitrogen concentration, Na, were nonlinearly related to , and photosynthetic capacity was linearly related to Na. Photosynthetic light acclimation resulted mainly from changes in Ma and leaf nitrogen partitioning, and to a lesser extent from changes in Nm. This behavior contrasts with photosynthetic light acclimation observed in other tree species like walnut (Juglans regia L.) in which acclimation results primarily from changes in Ma.     

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。     

Plasticity in mesophyll volume fraction modulates light-acclimation in needle photosynthesis in two pines

Acclimation potential of needle photosynthetic capacity varies greatly among pine species, but the underlying chemical, anatomical and morphological controls are not entirely understood. We investigated the light-dependent variation in needle characteristics in individuals of Pinus patula Schlect. & Cham., which has 19-31-cm long pendulous needles, and individuals of P. radiata D. Don., which has shorter (8-17-cm-long) stiffer needles. Needle nitrogen and carbon contents, mesophyll and structural tissue volume fractions, needle dry mass per unit total area (M(A)) and its components, volume to total area ratio (V/A(T)) and needle density (D = M(A)/(V/A(T))), and maximum carboxylase activity of Rubisco (V(cmax)) and capacity of photosynthetic electron transport (J(max)) were investigated in relation to seasonal mean integrated irradiance (Q(int)). Increases in Q(int) from canopy bottom to top resulted in proportional increases in both needle thickness and width such that needle total to projected surface area ratio, characterizing the efficiency of light interception, was independent of Q(int). Increased light availability also led to larger M(A) and nitrogen content per unit area (N(A)). Light-dependent modifications in M(A) resulted from increases in both V/A(T) and D, whereas N(A) changed because of increases in both M(A) and mass-based nitrogen content (N(M)) (N(A) = N(M)M(A)). Overall, the volume fraction of mesophyll cells increased with increasing irradiance and V/A(T) as the fraction of hypodermis and epidermis decreased with increasing needle thickness. Increases in M(A) and N(A) resulted in enhanced J(max) and V(cmax) per unit area in both species, but mass-based photosynthetic capacity increased only in P. patula. In addition, J(max) and V(cmax) showed greater plasticity in response to light in P. patula. Species differences in mesophyll volume fraction explained most of the variation in mass-based needle photosynthetic capacity between species, demonstrating that differences in plastic adjustments in mass-based photosynthetic activities among these representative individuals were mainly associated with contrasting investments in mesophyll cells. Greater area-based photosynthetic plasticity in P. patula relative to P. radiata was associated with larger increases in M(A) and mesophyll volume fraction with increasing irradiance. These data collectively demonstrate that light-dependent increases in mass-based nitrogen contents and photosynthetic activities were associated with an increased mesophyll volume fraction in needles at higher irradiances. They also emphasize the importance of light-dependent anatomical modifications in determining needle photosynthetic capacity.     

Does ontogeny modulate irradiance-elicited plasticity of leaf traits in saplings of rain-forest tree species? A test with Dicorynia guianensis and Tachigali melinonii (Fabaceae, Caesalpinioideae)

? Irradiance elicits a large plasticity in leaf traits, but little is known about the modulation of this plasticity by ontogeny. Interactive effects of relative irradiance and ontogeny were assessed on leaf traits for two tropical rainforest tree species: Dicorynia guianensis Amshoff and Tachigali melinonii (Harms) Barneby (Fabaceae, Caesalpinioideae).

? Eleven morphological and physiological leaf traits, relative to photosynthetic performance, were measured on saplings at three different architectural development stages (ASD 1, 2 and 3) and used to derive composite traits like photosynthetic N-use efficiency. Measurements were made along a natural irradiance gradient.

? The effect of ASD was very visible and differed between the two species. For Dicorynia guianensis, only leaf mass-per-area (LMA) significantly increased with ASDs whereas for Tachigali melinonii, almost all traits were affected by ASD: LMA, leaf N content and photosynthetic capacity increased from ASD 1 to ASD 3. Photosynthetic N-use-efficiency was not affected by ASD in any species.

? Leaf traits were severely modulated by irradiance, whereas the degree of plasticity was very similar among ASDs. Only few interactions were detected between irradiance and ASD, for leaf thickness, carbon content, and the ratio Chl/N in T. melinonii and for photosynthetic capacity in D. guianensis.

? We conclude that ontogenic development and irradiance-elicited plasticity modulated leaf traits, with almost no interaction, i.e., the degree of irradiance-elicited plasticity was stable across development stages and independent of ontogeny in these two species, at least in the early stages of development assessed here.

     

An analysis of light effects on foliar morphology,physiology, and light interception in temperate deciduous woody species of contrasting shade tolerance

Maximum Rubisco activities (V(cmax)), rates of photosynthetic electron transport (J(max)), and leaf nitrogen and chlorophyll concentrations were studied along a light gradient in the canopies of four temperate deciduous species differing in shade tolerance according to the ranking: Populus tremula L. < Fraxinus excelsior L. < Tilia cordata Mill. = Corylus avellana L. Long-term light environment at the canopy sampling locations was characterized by the fractional penetration of irradiance in the photosynthetically active spectral region (I(sum)). We used a process-based model to distinguish among photosynthesis limitations resulting from variability in fractional nitrogen investments in Rubisco (P(R)), bioenergetics (P(B), N in rate-limiting proteins of photosynthetic electron transport) and light harvesting machinery (P(L), N in chlorophyll and thylakoid chlorophyll-protein complexes). On an area basis, V(cmax) and J(max) (V(a) (cmax) and J(a) (max)) increased with increasing growth irradiance in all species, and the span of variation within species ranged from two (T. cordata) to ten times (C. avellana). Examination of mass-based V(cmax) and J(max) (V(m) (cmax) and J(m) (max)) demonstrated that the positive relationships between area-based quantities and relative irradiance mostly resulted from the scaling of leaf dry mass per area (M(A)) with irradiance. Although V(m) (cmax) and J(m) (max) were positively related to growth irradiance in C. avellana, and J(m) (max) was positively related to irradiance in P. tremula, the variation range was only a factor of two. Moreover, V(m) (cmax) and J(m) (max) were negatively correlated with relative irradiance in T. cordata. Rubisco activity in crude leaf extracts generally paralleled the gas-exchange data, but it was independent of light in T. cordata, suggesting that declining V(m) (cmax) with increasing relative irradiance was related to increasing diffusive resistances from the intercellular air spaces to the sites of carboxylation in this species. Because irradiance had little effect on foliar nitrogen concentration, the relationships of P(B) and P(R) with irradiance were similar to those of V(m) (cmax) and J(m) (max). Shade-intolerant species tended to have greater P(B) and P(R) and also larger V(a) (cmax) and J(a) (max) than more shade-tolerant species. However, for the whole material, P(B) and P(R) varied only about 50%, whereas V(a) (cmax) and J(a) (max) varied more than 15-fold, further emphasizing the importance of leaf anatomical plasticity in determining photosynthetic acclimation to high irradiance. Leaf chlorophyll concentrations and fractional nitrogen investments in light harvesting increased hyperbolically with decreasing irradiance to improve quantum use efficiency for incident irradiance. The effect of irradiance on P(L) was of the same order as its effect in the opposite direction on M(A), leading to either a constant model estimate of leaf absorptance with I(sum) or a slightly positive correlation. We conclude that leaf morphological plasticity is a more relevant determinant of foliage adaptation to high irradiance than foliage biochemical properties, whereas biochemical adaptation to low irradiance is of the same magnitude as the anatomical adjustments. Although shade-tolerant species did not have greater chlorophyll concentrations and P(L) than shade-intolerant species, they possessed lower M(A), and could maintain a more extensive foliar display for light capture with constant biomass investment in leaves.     

Ecophysiological responses of black walnut (Juglans nigra) to plantation thinning along a vertical canopy gradient

Ecophysiology of black walnut (Juglans nigra L.) along a vertical canopy gradient was studied in a mixed species plantation for the first (2007) and second (2008) growing seasons after thinning to understand response mechanisms to increased resource availability. Parameters were affected by the thinning treatment in 2008 only. Thinned trees showed increased light-saturated maximum photosynthesis (A_max) from 2007 to 2008 compared to non-thinned trees. This response was likely due to increased light levels near thinned trees (vs. changes in water or nutrient status), because thinning did not affect midday leaf water potential (Ψ_md), average daily soil water content (SWC), or leaf nitrogen content per unit area (N_a). Plantation thinning did not increase relative diameter growth during the experimental period. This may be due to low thinning intensity and mortality prior to thinning that reduced competition from first-tier neighbors. Certain leaf traits such as leaf mass per unit area (LMA) and N_a increased from the bottom to the upper canopy position, but did not influence thinning responses. Distribution patterns of photosynthetic parameters through the vertical canopy gradient were less defined than leaf structural traits such as LMA and N_a. Findings reflect black walnut's large variability in response to thinning.     

Growth and photosynthetic responses of Gallesia integrifolia (Spreng.) Harms and Schinus terebinthifolius Raddi seedlings in dense shade

We analyzed the growth and photosynthetic behavior of Gallesia integrifolia (‘pau-d’alho’) and Schinus terebinthifolius (‘aroeirinha’) under shade, seeking to obtain ecophysiological information for introducing seedlings of those species in previously established cacao agroforestry systems. Considering that light intensity under the shade of cacao trees varied between 5 and 10% daylight, 5 months old seedlings were exposed to four irradiance levels (25, 17, 10 and 5% daylight) for 92 days. With shade increase both species displayed trends of decrease leaf mass per unit leaf area, leaf area per plant (LA), relative growth rate (RGR) and net assimilation rate (NAR), and increase leaf area ratio (LAR). The mean values of light-saturated net photosynthetic rate (P _nmax) in 25 and 5% daylight were 12.8 and 8.0 μmol CO₂ m⁻² s⁻¹ for G. integrifolia and 17.9 and 7.4 μmol CO₂ m⁻² s⁻¹, respectively, for S. terebinthifolius. Based on the measurements of photosynthetic photon flux density and estimated values of photosynthetic saturated irradiance (Is) we concluded that, in all shaded conditions, the leaves of both species were under sub optimal light conditions to reach P _nmax. In spite of the lowest P _nmax values, RGR and NAR were significantly higher for G. integrifolia in all irradiance levels. Differences in growth rates can be explained by the higher values of LA, LAR and leaf mass ratio (LMR), as well as by the lower values of Is, photosynthetic compensation irradiance and dark respiration rates observed for G. integrifolia. Even though seedlings of G. integrifolia presented higher capacity to adapt under conditions of dense shade, we concluded that both species were under stress conditions induced by shade in light environments below 25% daylight. On a practical point of view it is possible to conclude that seedlings of both species should be introduced in light gaps, formed after the fall of big trees, or in places in which cacao trees are cultivated using large plant spacing.     

Between and within-site comparisons of structural and physiological characteristics and foliar nutrient content of 14 tree species at a wet,fertile site and a dry,infertile site in Panama

Structural and physiological characteristics and foliar nutrient content of 14 tree species were evaluated at two sites, one being seasonally wet with relatively fertile soils and the other being seasonally dry with relatively infertile soils. Differences in environmental stress between these sites drove the resulting differences in structural and physiological characteristics and leaf nutrient content of the investigated tree species. At the wet site, trees were more productive as site conditions allowed for greater photosynthetic activity to occur. The growth of pioneer tree species such as Spondias mombin, Guazuma ulmifolia, and Luehea seemanni, correlated strongly with high water-use efficiency and large, low-density leaves. Tree species, especially N-fixing species such as Albizia adinocephala, Albizia guachapele, Enterolobium cyclocarpum, and Gliricidia sepium, adapted to the greater levels of environmental stress at the dry site with infertile soils by increasing their water-use efficiency. Species differences were also significant, indicating that certain species adapted physiologically and structurally to environmental stress. Tree productivity operated under different structural and physiological constraints at each site. Leaf mass area (LMA), foliar N, and leaf area index (LAI) best predicted mass-based net photosynthetic capacity at the more fertile, wet site while foliar N was the best predictor of mass-based net photosynthetic capacity at the less fertile, dry site. Results from this study suggest the use of pioneer species at wet, fertile sites and N-fixing species at dry, infertile sites for restoration projects.     

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.     

Internal leaf anatomy and photosynthetic resource-use efficiency: interspecific and intraspecific comparisons

Leaf mass per unit area (LMA) and internal leaf anatomy often affect net gas exchange because of their effects on internal CO2 conductance to the site of carboxylation, internal shading, competition for CO2 among carboxylation sites, nitrogen concentration and its partitioning. To evaluate effects of LMA and leaf anatomy on CO2 assimilation, water-use efficiency (WUE) and nitrogen-use efficiency (NUE), we measured LMA, leaf thickness, the thickness of mesophyll components, and gas exchange rates at ambient CO2 concentration in leaves of six woody deciduous and evergreen species with different leaf life spans. In two species, CO2 assimilation was also estimated at saturating CO2 concentrations. There were interspecific differences in all morphological variables studied. Long-lived leaves had higher LMA and were thicker than short-lived leaves. Species with high LMA had low assimilation rates and NUE, both in ambient and saturating CO2 concentrations. Thus, in species with high LMA, assimilation was reduced by non-stomatal limitations, possibly because of a lower allocation of N to the photosynthetic machinery than in species with low LMA. Within a species, thicker leaves tended to have a lower tissue density. In intraspecific comparisons under field conditions, increasing internal air volume had positive effects on WUE, probably because of enhanced internal CO2 conductance to the site of carboxylation. We conclude that, in interspecific comparisons, different patterns of N partitioning strongly influence NUE, whereas in intraspecific comparisons, internal leaf anatomy is a key factor regulating resource-use efficiency.     

Foliar morphological and physiological plasticity in Picea abies and Abies alba saplings along a natural light gradient

The role of morphological versus physiological foliar plasticity in the capacity for, and mechanisms of, photosynthetic acclimation was assessed in Picea abies (L.) Karst. and Abies alba Mill. saplings in a forest gap-understory light gradient (relative irradiance, RI, ranging from 0.02 to 0.32). The species investigated showed a similar foliar morphological plasticity along the light gradient, at both the needle level (through alteration in leaf dry mass per area) and the shoot level (through alteration in the silhouette area ratio, e.g., shoot silhouette to projected needle area ratio). In both species chlorophyll (Chl) concentration on a mass basis decreased at increasing RI, but was independent of RI when expressed on an area basis. In contrast, leaf N concentration on a mass basis was independent of RI, but was positively influenced by RI when expressed on an area basis. The parameters describing photosynthetic performance at low light (dark respiration rate, apparent quantum yield and light compensation point) suggest that Abies alba was better suited to maintain a positive carbon balance in shaded conditions. By contrast, parameters describing biochemical capacity at high light (maximum electron transport rate, Jmax and maximum ribulose-1,5-biphosphate carboxylation capacity, Vcmax) indicate that only Picea abies was capable of acclimating physiologically to high photosynthetic photon flux densities (PPFDs) by increasing nitrogen partitioning to Rubisco and Vcmax/mass by increasing RI. These results support the hypothesis that interspecific differences in nitrogen partitioning within the photosynthetic apparatus may provide a mechanistic basis for species separation along a light gradient. The differences in photosynthetic plasticity observed are likely to influence regeneration patterns and habitat breadth of the species investigated. The limited ability of Abies alba saplings to exploit high-light conditions may be a competitive disadvantage in large canopy gaps and thus limit recruitment of this species to small gaps.     

Effects of water stress on irradiance acclimation of leaf traits in almond trees

Photosynthetic acclimation to highly variable local irradiance within the tree crown plays a primary role in determining tree carbon uptake. This study explores the plasticity of leaf structural and physiological traits in response to the interactive effects of ontogeny, water stress and irradiance in adult almond trees that have been subjected to three water regimes (full irrigation, deficit irrigation and rain-fed) for a 3-year period (2006-08) in a semiarid climate. Leaf structural (dry mass per unit area, N and chlorophyll content) and photosynthetic (maximum net CO(2) assimilation, A(max), maximum stomatal conductance, g(s,max), and mesophyll conductance, g(m)) traits and stem-to-leaf hydraulic conductance (K(s-l)) were determined throughout the 2008 growing season in leaves of outer south-facing (S-leaves) and inner northwest-facing (NW-leaves) shoots. Leaf plasticity was quantified by means of an exposure adjustment coefficient (ε=1-X(NW)/X(S)) for each trait (X) of S- and NW-leaves. Photosynthetic traits and K(s-l) exhibited higher irradiance-elicited plasticity (higher ε) than structural traits in all treatments, with the highest and lowest plasticity being observed in the fully irrigated and rain-fed trees, respectively. Our results suggest that water stress modulates the irradiance-elicited plasticity of almond leaves through changes in crown architecture. Such changes lead to a more even distribution of within-crown irradiance, and hence of the photosynthetic capacity, as water stress intensifies. Ontogeny drove seasonal changes only in the ε of area- and mass-based N content and mass-based chlorophyll content, while no leaf age-dependent effect was observed on ε as regards the physiological traits. Our results also indicate that the irradiance-elicited plasticity of A(max) is mainly driven by changes in leaf dry mass per unit area, in g(m) and, most likely, in the partitioning of the leaf N content.     

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.