Determinants of biomass production in hybrid willows and prediction of field performance from pot studies

Pot experiments are often performed to assess plant physiological traits and relationships among growth traits under controlled environments. However, the reliability of pot studies for predicting the growth and performance of trees in the field has rarely been rigorously assessed. We evaluated the suitability of pot experiments for predicting field performance, measured as shoot biomass production, by investigating determinants of growth in hybrid willows (Salix spp.) grown under various environmental conditions in the field, and by comparing the data with the results from a corresponding pot study. Biomass production in six hybrid willow clones, bred for use as bio-fuels, was assessed in three field trials located in central and southeastern Sweden throughout the first 3-year cutting cycle. The determinants of biomass productivity, measured as biomass allocation and nitrogen (N) economy, were identified in one of the field trials. Key traits for shoot biomass production in the field were total leaf area and total amount of N; plant N losses by shed leaves were only partly controlled by leaf-litter N concentration. These key traits were also obtained from the pot study and related to shoot biomass production and abscission-leaf N loss in the field. Total leaf area and total N pool of plants grown in pot experiments were good predictors of long-term biomass production in the field, whereas shoot biomass production, specific leaf area and tissue N concentration of pot-grown plants were less suitable as predictors of field performance. Relationships between the key traits and shoot biomass production were clone-specific, indicating the need for analysis of growth traits at the clone level if field performance of trees is to be evaluated based on data from pot studies. Nutrient loss components are important for tree performance in the long term and evaluations of nutrient loss characteristics at the individual-tree level should focus on nutrient pools lost rather than on nutrient concentrations in abscised plant parts.     

Co-optimal distribution of leaf nitrogen and hydraulic conductance in plant canopies

Leaf properties vary significantly within plant canopies, due to the strong gradient in light availability through the canopy, and the need for plants to use resources efficiently. At high light, photosynthesis is maximized when leaves have a high nitrogen content and water supply, whereas at low light leaves have a lower requirement for both nitrogen and water. Studies of the distribution of leaf nitrogen (N) within canopies have shown that, if water supply is ignored, the optimal distribution is that where N is proportional to light, but that the gradient of N in real canopies is shallower than the optimal distribution. We extend this work by considering the optimal co-allocation of nitrogen and water supply within plant canopies. We developed a simple 'toy' two-leaf canopy model and optimized the distribution of N and hydraulic conductance (K) between the two leaves. We asked whether hydraulic constraints to water supply can explain shallow N gradients in canopies. We found that the optimal N distribution within plant canopies is proportional to the light distribution only if hydraulic conductance, K, is also optimally distributed. The optimal distribution of K is that where K and N are both proportional to incident light, such that optimal K is highest to the upper canopy. If the plant is constrained in its ability to construct higher K to sun-exposed leaves, the optimal N distribution does not follow the gradient in light within canopies, but instead follows a shallower gradient. We therefore hypothesize that measured deviations from the predicted optimal distribution of N could be explained by constraints on the distribution of K within canopies. Further empirical research is required on the extent to which plants can construct optimal K distributions, and whether shallow within-canopy N distributions can be explained by sub-optimal K distributions.     

Distribution of leaf mass per unit area and leaf nitrogen concentration determine partitioning of leaf nitrogen within tree canopies

Distribution of leaf nitrogen with respect to leaf mass per unit area (M(a)), nitrogen per unit mass (N(m)) and nitrogen per unit area (N(a)) within peach (Prunus persica L.) tree canopies was studied in two field experiments. In one experiment, leaf light exposure and M(a) were measured on leaves from different canopy positions of peach trees subjected to five nitrogen (N) fertilization treatments. Leaf light exposure and M(a) were linearly related and the relationship was independent of N fertilization. In a subsequent experiment, N fertilizer was applied to previously unfertilized trees in midsummer, after shoot growth had terminated. Application of N fertilizer did not affect mean canopy M(a). Fertilization increased N(m) of all leaves throughout the canopy compared with non-fertilized trees. No significant relationship between N(m) and M(a) was found in either fertilized or control trees. There was a linear relationship between N(a) and M(a) and the slope of the relationship was increased by N fertilizer application. We conclude that distribution of N(a) in peach tree canopies is primarily a function of M(a) partitioning with light and N(m), which is related to soil N availability.     

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

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

Leaf traits in relation to crown development, light interception and growth of elite families of loblolly and slash pine

Crown architecture and size influence leaf area distribution within tree crowns and have large effects on the light environment in forest canopies. The use of selected genotypes in combination with silvicultural treatments that optimize site conditions in forest plantations provide both a challenge and an opportunity to study the biological and environmental determinants of forest growth. We investigated tree growth, crown development and leaf traits of two elite families of loblolly pine (Pinus taeda L.) and one family of slash pine (P. elliottii Mill.) at canopy closure. Two contrasting silvicultural treatments -- repeated fertilization and control of competing vegetation (MI treatment), and a single fertilization and control of competing vegetation treatment (C treatment) -- were applied at two experimental sites in the West Gulf Coastal Plain in Texas and Louisiana. At a common tree size (diameter at breast height), loblolly pine trees had longer and wider crowns, and at the plot-level, intercepted a greater fraction of photosynthetic photon flux than slash pine trees. Leaf-level, light-saturated assimilation rates (A(max)) and both mass- and area-based leaf nitrogen (N) decreased, and specific leaf area (SLA) increased with increasing canopy depth. Leaf-trait gradients were steeper in crowns of loblolly pine trees than of slash pine trees for SLA and leaf N, but not for A(max). There were no species differences in A(max), except in mass-based photosynthesis in upper crowns, but the effect of silvicultural treatment on A(max) differed between sites. Across all crown positions, A(max) was correlated with leaf N, but the relationship differed between sites and treatments. Observed patterns of variation in leaf properties within crowns reflected acclimation to developing light gradients in stands with closing canopies. Tree growth was not directly related to A(max), but there was a strong correlation between tree growth and plot-level light interception in both species. Growth efficiency was unaffected by silvicultural treatment. Thus, when coupled with leaf area and light interception at the crown and canopy levels, A(max) provides insight into family and silvicultural effects on tree growth.     

Stoichiometry of foliar carbon constituents varies along light gradients in temperate woody canopies: implications for foliage morphological plasticity

Foliar morphology and chemical composition were examined along a light gradient in the canopies of five deciduous temperate woody species, ranked according to shade-tolerance as Populus tremula L. < Fraxinus excelsior L. < Tilia cordata Mill. = Corylus avellana L. < Fagus sylvatica L. Foliar carbon was divided between structural (cell-wall polysaccharides, lignin) and nonstructural (proteins, ethanol-soluble carbohydrates, starch) fractions. Foliar morphology of all species was strongly affected by irradiance. Both leaf dry mass per area (M(A)), a product of leaf density and thickness, and leaf dry to fresh mass ratio (D(w)), characterizing the apoplastic leaf fraction, increased with increasing relative irradiance (I(sum), calculated as the weighted mean of fractional penetration of diffuse and direct irradiance). Though the relationships were qualitatively identical among the taxa, more shade-tolerant species generally had lower values of M(A) than shade-intolerant species, and their morphological relationships with irradiance were curvilinear; however, there were no signs of saturation even at the highest irradiances in shade-intolerant species. In all species, lignin concentrations increased and cell-wall polysaccharide concentrations decreased with increasing irradiance. Consequently, biomass investment in structural leaf components appeared to be relatively constant along light gradients. The relationship between irradiance and structural compounds tended to be asymptotic in the more shade-tolerant species, whereas M(A) was linearly correlated with concentrations of structural leaf components, suggesting that similar factors were responsible for the curvature in the morphological and chemical relationships with irradiance. Because lignin increases tissue elastic modulus thereby rendering leaves more resistant to low leaf water potentials, we conclude that changes in stoichiometry of cell wall components were related to foliage acclimation to the gradients of water deficit that develop in the canopy and inherently accompany light gradients. We also conclude that increased lignification decreased leaf expansion growth, and that species differences in lignification were partly responsible for the observed interspecific variability in morphological plasticity. Analysis of structural leaf compounds provided no indication of how shade-intolerant species with low investments in lignin acclimated to gradients of water availability in the canopy. Because shade-intolerant species generally had higher capacities for photosynthesis than shade-tolerant species, we postulated that they should also have a greater ability for osmotic adjustment of leaf water potential with photosynthates. The concentrations of soluble carbohydrates increased with increasing irradiance in all species; however, the osmotic adjustment achieved in this way was similar in all species, except for shade-intolerant F. excelsior, which had a lower potential for osmotic adjustment with carbohydrates than the other taxa. Although we did not determine whether the gradients of stem water potential and leaf water deficits were similar in canopies of different species, we demonstrated that water relations play a central role in determining foliar structure and composition along light gradients in the canopy.     

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.     

Influence of foliar phenology and shoot inclination on annual photosynthetic gain in individual beech saplings: A functional–structural modeling approach

We developed a functional–structural plant model for Fagus crenata saplings and calculated annual photosynthetic gains to determine the influences of foliar phenology and shoot inclination on the carbon economy of saplings. The model regenerated the three-dimensional shoot structure and spatial and temporal display of leaves; we calculated the hourly light interception of each leaf with a detailed light model that allowed us to estimate hourly leaf photosynthetic gain taking leaf age into account. To evaluate the importance of simultaneous foliar phenology and slanting shoots in beech saplings, we calculated the photosynthetic budgets for saplings with contrasting foliar phenologies and shoot inclinations. In our simulations, we distinguished between simultaneous and successive foliar phenologies, upright and slanting shoot inclinations, and environments with and without a vertical gradient in light intensity. Other model parameters (including photosynthesis vs. light curve, leaf size, and leaf shape) were obtained directly from live beech saplings. With no vertical gradient in light intensity, modeled saplings with simultaneous foliar phenology and slanting shoots (as in live beech) had larger annual photosynthetic gains than saplings with other combinations of traits. Hence, simultaneous foliar phenology and slanting shoots are efficient ways to display leaves in the shaded forest understory light regime where beech saplings thrive. In the presence of vertical light gradients, which can occur in canopy gaps, saplings with upright shoots had larger annual photosynthetic gains than counterparts with slanting shoots. Although mean daily photosynthetic gains of saplings with successive foliar phenology were elevated by exposing leaves to strong light when young and productive, the annual photosynthetic budget of these saplings was reduced (compared to saplings with simultaneous foliar phenology) by their relatively short leaf lifespan. Overall, our results suggest that slanting shoots with simultaneous foliar phenology are particularly successful in shaded environments, where beech often dominates, because they appear to maximize the annual carbon budget by avoiding self-shading and extending leaf lifespans.     

Shoot structure and growth along a vertical profile within a Populus-Tilia canopy

We investigated shoot growth patterns and their relationship to the canopy radiation environment and the distribution of leaf photosynthetic production in a 27-m-tall stand of light-demanding Populus tremula L. and shade-tolerant Tilia cordata Mill. The species formed two distinct layers in the leaf canopy and showed different responses in branch architecture to the canopy light gradient. In P. tremula, shoot bifurcation decreased rapidly with decreasing light, and leaf display allowed capture of multidirectional light. In contrast, leaf display in T. cordata was limited to efficient interception of unidirectional light, and shoot growth and branching pattern facilitated relatively rapid expansion into potentially unoccupied space even in the low light of the lower canopy. At the canopy level, T. cordata had higher photosynthetic light-use efficiency than P. tremula, whereas P. tremula had higher nitrogen-use efficiency than T. cordata. However, at the individual leaf level, both species had similar efficiencies under comparable light conditions. Production of new leaf area in the canopy followed the pattern of photosynthetic production. However, the species differed substantially in extension growth and space-filling strategy. Light-demanding P. tremula expanded into new space with a few long shoots, with shoot length strongly dependent on photosynthetic photon flux density (PPFD). Production of new leaf area and extension growth were largely uncoupled in this species because short shoots, which do not contribute to extension growth, produced many new leaves. Thus, in P. tremula, the growth pattern was strongly directed toward the top of the canopy. In contrast, in shade-tolerant T. cordata, shoot growth was weakly related to PPFD and more was invested in long shoot growth on a leaf area basis compared with P. tremula. However, this extension growth was not directed and may serve as a passive means of avoiding self-shading. This study supports the hypothesis that, for a particular species, allocation patterns and crown architecture contribute as much to shade tolerance as leaf-level photosynthetic acclimation.     

Spatial distribution of leaf morphological and physiological characteristics in relation to local radiation regime within the canopies of 3-year-old Populus clones in coppice culture

Spatial distributions of leaf characteristics relevant to photosynthesis were compared within high-density coppice canopies of Populus spp. of contrasting genetic origin. We studied three clones representative of the range in growth potential, leaf morphology, coppice and canopy structure: Clone Hoogvorst (Hoo) (Populus trichocarpa Torr. & Gray x Populus deltoides Bartr. & Marsh), Clone Fritzi Pauley (Fri) (Populus trichocarpa Torr. & Gray) and Clone Wolterson (Wol) (Populus nigra L.). Leaf area index ranged from 2.7 (Fri and Wol) to 3.8 (Hoo). The clones exhibited large vertical variation in leaf area density (0.02-1.42 m2 m-3). Leaf dry mass per unit leaf area (DM(A)) increased with increasing light in Clones Hoo and Fri, from about 56 g m-2 at the bottom of the canopy to 162 g m-2 at the top. In Clone Wol, DM(A) varied only from 65 to 100 g m-2, with no consistent relationship with respect to light. Conversely, nitrogen concentration on a mass basis was nearly constant (around 1.3-2.1%) within the canopies of Clones Hoo and Fri, but increased strongly with light in Clone Wol, from 1.4% at the bottom of the canopy to 4.1% at the top. As a result, nitrogen per unit leaf area (N(A)) increased with light in the canopies of all clones, from 0.9 g m-2 at the bottom to 2.9 g m-2 at the top. Although a single linear relationship described the dependence of maximum carboxylation rate (17-93 micromol CO2 m-2 s-1) or electron transport capacity (45-186 micromol electrons m-2 s-1) on N(A), for all clones, Clone Wol differed from Clones Hoo and Fri by exhibiting a higher dark respiration rate at low N(A) (1.8 versus 0.8 micromol CO2 m-2 s-1).     

不同密度杉木林内辐射与叶面积垂直分布对生长的影响

本文通过对八年生杉木人工林内太阳辐射与叶面积垂直分布的观测与研究，用计算机绘图与计算，得出以下结果：杉木叶面积的垂直分布和叶面积指数与密度有关，冠形因密度增加由圆锥形变为圆柱形，叶面积指数是先上升后下降。杉木林内辐射的消减随密度的增加而加剧，在冠层深3／4处趋于平缓，当郁闭度低于0．85时，林冠下辐射有回升，用累加叶面积指数分层计算林冠消光系数，可减少因叶片分布不均匀而产生的误差，用累计值计算辐射，可简化观测与计算，用辐射吸收率、叶面积指数，杉木个体生物量及年均生物增长量4个指标对杉木林生长作综合评价，八年生杉木林生长的较适宜工是2m×1m。     

Effects of shade on growth, biomass allocation and leaf morphology in European yew (Taxus baccata L.)

The impact of shade on the growth of European yew (Taxus baccata L.) saplings was investigated over a three-year period using artificial shading to simulate four different light regimes (3, 7, 27 and 100 % relative photosynthetic photon flux density, RPPFD). There was no mortality attributable to shading even under the 3 % RPPFD treatment. Increasing shade was positively associated with specific leaf area, leaf length, leaf width and total chlorophyll content, but negatively associated with plant height, stem diameter, total dry weight and root to leaf and shoot ratio. Discoloration of the foliage occurred in plants grown in 100 % RPPFD conditions (resulting in reduced growth rates) and those transferred to 100 % RPPFD conditions after being shade-acclimated for 2 years. Evidence suggests that T. baccata has the ability to regenerate beneath a lighter canopy but beneath denser canopies gap dynamics will play an important role in facilitating successful regeneration and this needs to be reflected in management of natural populations of this declining species.     

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.     

Analysis of the sensitivity of absorbed light and incident light profile to various canopy architecture and stand conditions

We analyzed the effect of simplifying assumptions in canopy representation of radiation transfer models, comparing modeled diffuse non-interceptance and photosynthetic photon flux density with measurements at different layers of complex pine-broadleaved canopy with large seasonal variation of leaf area index. The most detailed model included clumping of trees (i.e.,?stand density) and a vertical specification of leaf angle distribution and shoot clumping. A less detailed model replaced the vertically specified variables with their means. The most parsimonious model accounted for neither shoot clumping nor stand density. The vertical specification of shoot clumping and leaf angle distribution only slightly improved vertical and seasonal openness and light estimates over using mean values. Further simplification had little effect on total absorbed light but was more risky for estimates of the vertical distributions of openness and light absorbed by the canopy, which will affect photosynthesis estimates due to the non-linearity of photosynthetic light response. Including woody surfaces in winter, when leaf area was low, was essential for reproducing the measurements correctly. A sensitivity analysis showed that ignoring (i)?shoot clumping could result in a substantial overestimation of total absorbed light with errors increasing with decreasing leaf area and (ii) stand density in sparse stands could lead to substantial overestimation of total absorbed light, and the effect is largely independent of leaf area. Also, (iii) the effect of changing leaf angle distribution increased with decreasing leaf area, and was larger and more persistent along the leaf area range with increasing shoot clumping. Overall, accounting for the effect of tree clumping on absorbed light is most important in stands composed of species where leaves are not very clumped (e.g., broadleaved). However, even in forests with highly clumped shoots (i.e., coniferous), an accurate estimation of absorbed light distribution in stands requires incorporation of stand density in the model.     

Morphological acclimation and growth of ash (Fraxinus pennsylvanica Marsh.) advance regeneration following overstory harvesting in a Mississippi River floodplain forest

Stand-level growth responses and plant-level patterns of biomass accumulation and distribution were examined to learn how stand structure influences morphological acclimation and growth of green ash (Fraxinus pennsylvanica Marsh.) advance regeneration following overstory harvesting. Nine, 20-ha plots that received clearcut harvesting (100% basal area removal), partial harvesting (50% basal area removal), or no harvesting (control) were sampled to measure height, root-collar diameter, leaf, stem and root biomass, and leaf mass ratio (LMR), stem mass ratio (SMR) and root mass ratio (RMR) of ash regeneration. Six years after treatment, plot-level analyses indicated that ash growth was greatest in plots receiving clearcut harvesting, and least in control plots. Examination of LMR, SMR and RMR revealed that this growth response was not associated with acclimation that altered plant morphology. Total biomass ranged 275-fold among sampled plants, and much of this variation was accounted for by measurements of stand leaf area index (LAI). Along the gradient of stand LAI, values greater than 2 inhibited biomass accumulation. Stand LAI values less than 1.5 promoted ash biomass accumulation which reached a maximum where LAI values approached 0.7 and tapered above or below this value. Our findings indicate that green ash regeneration can be managed beneath light canopy cover, and the ability of seedlings to establish and persist beneath closed canopies and vigorously respond to release without having to endure prolonged morphological acclimation provides flexibility in developing regeneration protocols.     

Summary models for light interception and light-use efficiency of non-homogeneous canopies

The application of detailed models of canopy photosynthesis rely on the estimation of attenuation of light in the canopy. This attenuation is readily estimated with the Lambert-Beer law when the canopy is homogeneous. In reality, forest canopies are far from homogeneous, and this has led to the use of detailed light extinction models that account for grouping of foliage between and within trees. Because such models require detailed parameterization and fine resolution inputs, they are impractical in larger-scale applications. Thus, there is interest in simplified models that can be readily parameterized. We developed two equations that can be used to estimate mean annual light interception by single unshaded trees and by stands of Poisson distributed trees. Interception by single trees is a function of crown surface area, the ratio of leaf area to crown surface area, the extinction coefficient in a homogeneous canopy--which can be determined separately--and one empirical parameter that depends on the mean solar angle. The summary model was tested against a detailed model of interception, and showed good agreement, although with slight bias. The results showed that crown surface area is a good summary variable for crown size and shape, because errors are independent of crown shape (ellipsoids, cones and height:width ratios). We also tested whether canopy photosynthesis is proportional to light interception across canopies differing in structure and leaf area index, and found that light-use efficiency is influenced by canopy structure. The model is useful in larger-scale applications because it can be parameterized with available data without the need for additional empirical parameters. It can also be used to study the effect of stand structure on mean annual light interception and productivity.     

Influence of canopy light environment and nitrogen availability on leaf photosynthetic characteristics and photosynthetic nitrogen-use efficiency of field-grown nectarine trees

Relationships between CO(2) assimilation at light saturation (A(max)), nitrogen (N) content and weight per unit area (W(A)) were studied in leaves grown with contrasting irradiances (outer canopy versus inner canopy) and N supply rates in field-grown nectarine trees Prunus persica L. Batsch. cv. Fantasia. Both A(max) and N content per unit leaf area (N(A)) were linearly correlated to W(A), but leaves in the high-N treatment had higher N(A) and A(max) for the same value of W(A) than leaves in the low-N treatment. The curvilinear relationship between photosynthesis and total leaf N was independent of treatments, both when expressed per unit leaf area A(maxA) and N(A)) and per unit leaf weight (A(maxW) and N(W)), but the relationship was stronger when data were expressed on a leaf area basis. Both A(maxA) and N(A) were higher for outer canopy leaves than for inner canopy leaves and A(maxW) and N(W) were higher for leaves in the high-N treatment than for leaves in the low-N treatment. The relationship between A(max) and N resulted in a similar photosynthetic nitrogen-use efficiency at light saturation (A(max)NUE) for both N and light treatments. Photosynthetic nitrogen-use efficiency was similar among treatments throughout the whole light response curve of photosynthesis. Leaves developed in shade conditions did not show higher N-use efficiency at low irradiance. At any intercellular CO(2) partial pressure (C(i)), photosynthetic CO(2) response curves were higher for outer canopy leaves and, within each light treatment, were higher for the high-N treatments than for the low-N treatments. Consequently, most of the differences among treatments disappeared when photosynthesis was expressed per unit N. However, slightly higher assimilation rates per unit N were found for outer canopy leaves compared with inner canopy leaves, in both N treatments. Because higher daily irradiance within the canopies of the low-N trees more than compensated for the lower photosynthetic performances of these leaves compared to the leaves of high-N trees, daily carbon gain (and N-use efficiency on a daily assimilation basis) per leaf was higher for the low-N treatment than for the high-N treatment in both outer and inner canopy leaves.     

Interspecific and environmentally induced variation in foliar dark respiration among eighteen southeastern deciduous tree species

We measured variations in leaf dark respiration rate (Rd) and leaf nitrogen (N) across species, canopy light environment, and elevation for 18 co-occurring deciduous hardwood species in the southern Appalachian mountains of western North Carolina. Our overall objective was to estimate leaf respiration rates under typical conditions and to determine how they varied within and among species. Mean dark respiration rate at 20 degrees C (Rd,mass, micromol CO2 per kg leaf dry mass per s) for all 18 species was 7.31 micromol per kg per s. Mean Rd,mass of individual species varied from 5.17 micromol per kg per s for Quercus coccinea Muenchh. to 8.25 micromol per kg per s for Liriodendron tulipifera L. Dark respiration rate varied by leaf canopy position and was higher in leaves collected from high-light environments. When expressed on an area basis, dark respiration rate (Rd,area, micromol CO2 per kg leaf dry area per s) showed a strong linear relationship with the predictor variables leaf nitrogen (Narea, g N per square m leaf area) and leaf structure (LMA, g leaf dry mass per square m leaf area) (r squared = 0.62). This covariance was largely a result of changes in leaf structure with canopy position; smaller thicker leaves occur at upper canopy positions in high-light environments. Mass-based expression of leaf nitrogen and dark respiration rate showed that nitrogen concentration (Nmass, mg N per g leaf dry mass) was only moderately predictive of variation in Rd,mass for all leaves pooled (r squared = 0.11), within species, or among species. We found distinct elevational trends, with both Rd,mass and Nmass higher in trees originating from high-elevation, cooler growth environments. Consideration of interspecies differences, vertical gradients in canopy light environment, and elevation, may improve our ability to scale leaf respiration to the canopy in forest process models.     

Relative importance of photosynthetic physiology and biomass allocation for tree seedling growth across a broad light gradient

Studies of tree seedling physiology and growth under field conditions provide information on the mechanisms underlying inter- and intraspecific differences in growth and survival at a critical period during forest regeneration. I compared photosynthetic physiology, growth and biomass allocation in seedlings of three shade-tolerant tree species, Virola koschynii Warb., Dipteryx panamensis (Pittier) Record & Mell and Brosimum alicastrum Swartz., growing across a light gradient created by a forest-pasture edge (0.5 to 67% diffuse transmittance (%T)). Most growth and physiological traits showed nonlinear responses to light availability, with the greatest changes occurring between 0.5 and 20 %T. Specific leaf area (SLA) and nitrogen per unit leaf mass (N mass) decreased, maximum assimilation per unit leaf area (A area) and area-based leaf N concentration (N area) increased, and maximum assimilation per unit leaf mass (A mass) did not change with increasing irradiance. Plastic responses in SLA were important determinants of leaf N and A area across the gradient. Species differed in magnitude and plasticity of growth; B. alicastrum had the lowest relative growth rates (RGR) and low plasticity. Its final biomass varied only 10-fold across the light gradient. In contrast, the final biomass of D. panamensis and V. koschynii varied by 100- and 50-fold, respectively, and both had higher RGR than B. alicastrum. As light availability increased, all species decreased biomass allocation to leaf tissue (mass and area) and showed a trade-off between allocation to leaf area at a given plant mass (LAR) and net gain in mass per unit leaf area (net assimilation rate, NAR). This trade-off largely reflected declines in SLA with increasing light. Finally, A area was correlated with NAR and both were major determinants of intraspecific variation in RGR. These data indicate the importance of plasticity in photosynthetic physiology and allocation for variation in tree seedling growth among habitats that vary in light availability.     

Nitrogen availability, local light regime and leaf rank effects on the amount and sources of N allocated within the foliage of young walnut (Juglans nigra x regia) trees

Early season leaf growth depends largely on nitrogen (N) provided by remobilization from storage, and many studies have tested the effect of N availability to roots on the amount of N provided for new leaf development by remobilization. Although it is well known that the light regime experienced by a leaf influences the amount of N per unit leaf area (LA), the effect of the local light regime on the amount of N derived either directly from root uptake or from remobilization for early season leaf growth has never been tested at an intra- canopy scale. The objective of this study was to quantify the relative importance of (1) N availability to roots, (2) local light regime experienced by the foliage (at the shoot scale) and (3) leaf rank along the shoot, on the total amount of N allocated to leaves and on the proportions of N provided by remobilization and root uptake. To quantify the importance of N uptake and remobilization as sources of leaf N, potted hybrid walnut trees (Juglans nigra L. x regia L.) were grown outdoors in sand and fed with a labeled ((15)N) nutrient solution. By removing the apical bud, the trees were manipulated to produce only two shoots. The experimental design had two factors: (1) high (HN; 8 mol N m(-3)) and low (LN; 2 mol N m(-3)) N availability; and (2) high (HL; 90% of incident photosynthetically active photon flux (PPF)) and low (LL; 10% of incident PPF) light. Total leaf N per tree was unaffected by either N availability or irradiance. The HN treatment increased the amount of leaf N derived from root uptake at the whole-tree scale (typically around 8 and 2% in the HN and LN treatments, respectively). Nitrogen allocation within foliage of individual trees was controlled by the local light regime, which strongly affected individual leaf characteristics as leaf mass per unit LA and area- based amount of leaf (N(a)). Decreasing the light availability to a branch decreased the amount of N allocated to it, benefiting the less shaded branches. In contrast, shading of the lower branch did not affect the fraction of total leaf N remobilized for either the lower, shaded branch or the upper, unshaded branch. The relevance of these findings for tree growth modeling is discussed.