Biomass investment in leaf lamina versus lamina support in relation to growth irradiance and leaf size in temperate deciduous trees

Foliar biomass investment in support and assimilative compartments was studied in four temperate deciduous tree species along a natural light gradient across the canopy. The species ranked according to shade tolerance as Betula pendula Roth. < Populus tremula L. < Fraxinus excelsior L. < Tilia cordata Mill. Long-term light conditions at sampling locations were characterized as seasonal mean integrated quantum flux density (Q(int), mol m(-2) day(-1)) estimated by a method combining hemispherical photography and light measurements with quantum sensors. Leaf morphology was altered by Q(int) in all species. Both lamina and petiole dry mass per lamina area (LMA and PMA, respectively) increased with increasing Q(int). Shade-tolerant species had lower LMA at low Q(int) than shade-intolerant species; however, PMA was not related to shade tolerance. Across species, the ratio of petiole dry mass to lamina dry mass (PMR) varied from 0.07 to 0.21. It was independent of Q(int) in the simple-leaved species, but decreased with increasing Q(int) in the compound-leaved F. excelsior, which also had the largest foliar biomass investment in petioles. Differences in leaf mass and area, ranging over four orders of magnitude, provided an explanation for the interspecific variability in PMR. Species with large leaves also had greater biomass investments in foliar support than species with smaller leaves. This relationship was similar for both simple- and compound-leaved species. There was a negative relationship between PMR and petiole N concentration, suggesting that petioles had greater carbon assimilation rates and paid back a larger fraction of their construction cost in species with low PMR than in species with high PMR. This was probably the result of a negative relationship between PMR and petiole surface to volume ratio. Nevertheless, petioles had lower concentrations of mineral nutrients than laminas. Across species, the ratio of petiole N to lamina N varied from only 3 to 6%, demonstrating that petiole costs are less in terms of nutrients than in terms of total biomass, and that the petiole contribution to carbon assimilation is disproportionately lower than that of the lamina contribution.     

Differences in chemical composition relative to functional differentiation between petioles and laminas of Fraxinus excelsior

Differences in structural and nonstructural carbohydrates, lignin and chlorophyll, and Rubisco (ribulose-1,5-bisphosphate carboxylase/oxygenase) activity between petioles and leaflets were studied along a canopy light gradient in Fraxinus excelsior L., which has pinnate compound leaves and up to 20% of foliar biomass invested in petioles. Long-term light conditions at the sampling locations were characterized by values of seasonal mean integrated quantum flux density (Q(int), mol m(-2) day(-1)) estimated by combining data from hemispherical photographs at the sampling locations with measurements of global solar radiation above the canopy during the growing season. The contribution of petioles to leaf carbon assimilation was disproportionally lower than that of leaf laminas. Though the light relationships of assimilative compounds-foliar chlorophyll concentration increasing with decreasing Q(int) to improve leaf absorptance, foliar N concentration and Rubisco activity being relatively constant along the light gradient-were similar for both petioles and leaflets, petiole nitrogen and chlorophyll concentrations were only 30% and 10%, respectively, of those of leaflets. Nonstructural carbohydrate concentration was about 20% higher in petioles than in leaf laminas, indicating that petioles also serve as storage tissues for photosynthates. Relationships between foliar structural carbon components and irradiance-increasing lignin (L) and decreasing structural polysaccharide (SP) concentrations with increasing Q(int)-were qualitatively similar for petioles and leaflets. However, petioles had lower L, but higher SP and total investment in structural compounds (L + SP) than leaflets. Greater lignification at high irradiances in leaflets than in petioles was attributed to greater water stresses at high light, and to more variable water contents of actively transpiring leaflets. Low lignin concentration in combination with high osmotically active carbohydrate concentrations in petioles suggest that turgor plays an important role in the mechanical properties of petioles. As a result of lower lignin and protein concentrations, the glucose cost of petiole construction (g glucose per g dry mass) was about 5% lower than that of leaf laminas.     

Diversity of leaf traits related to productivity in 31 Populus deltoides x Populus nigra clones

To test if some leaf parameters are predictors of productivity in a range of Populus deltoides (Bartr.) Marsh. x P. nigra L. clones, we assessed leaf traits and productivity in 2-month-old rooted cuttings from 31 clones growing in 4-l pots in a greenhouse, under conditions of controlled temperature and optimal irrigation. We evaluated four groups of variables describing (1) productivity (total biomass), (2) leaf growth (total leaf number increment and total leaf area increment rate), (3) leaf structure (specific leaf area and nitrogen and carbon contents) and (4) carbon isotope discrimination (delta), which is negatively correlated with time-integrated water-use efficiency. High-yielding clones did not necessarily display high leaf growth rates, but they displayed a larger total leaf area, lower specific leaf area and lower leaf nitrogen concentration than clones with low productivity. Total leaf area was mainly controlled by maximal individual leaf area and total leaf area increment rate (r = 0.51 and 0.56, respectively). Carbon isotope discrimination did not correlate with total biomass, but it was associated with total number of leaves and total leaf area increment rate (r = 0.39 and 0.45, respectively). Therefore, leaf area and specific leaf area were better indicators of productivity than leaf growth traits. The observed independence of delta from biomass production provides opportunities for selecting poplar clones combining high productivity and high water-use efficiency.     

Differences in leaf functional traits between red and green leaves of two evergreen shrubs Photinia × fraseri and Osmanthus fragrans

Leaf functional traits are adaptations that enable plants to live under different environmental conditions. This study aims to evaluate the differences in leaf functional traits between red and green leaves of two evergreen shrubs Photinia 9 fraseri and Osmanthus fragrans. Specific areas of red leaves are higher than that of green leaves in both species. Thus, the material investment per unit area and per lamina of red leaves is significantly lower than that of green leaves, implying an utmost effort of red leaves to increase light capture and use efficiency because of their low leafchlorophyll concentration. The higher petiole length of green leaves compared with that of red leaves indicates that adult green leaves may have large fractional biomass allocation to support the lamina structures in capturing light with maximum efficiency and obtaining a high growth rate. The high range of the phenotypic plasticity of leaf size, leaf thickness,single-leaf wet and dry weights, and leaf moisture of green leaves may be beneficial in achieving efficient control of water loss and nutrient deprivation. The high range of phenotypic plasticity of leaf chlorophyll concentration of red leaves may be advantageous in increasing resource(especially light) capture and use efficiency because this leaf type is juvenile in the growth stage and has low leaf-chlorophyll concentration.     

Differences in leaf functional traits between red and green leaves of two evergreen shrubs <Emphasis Type="Italic">Photinia</Emphasis> × <Emphasis Type="Italic">fraseri</Emphasis> and <Emphasis Type="Italic">Osmanthus fragrans</Emphasis>

Leaf functional traits are adaptations that enable plants to live under different environmental conditions. This study aims to evaluate the differences in leaf functional traits between red and green leaves of two evergreen shrubs Photinia × fraseri and Osmanthus fragrans. Specific areas of red leaves are higher than that of green leaves in both species. Thus, the material investment per unit area and per lamina of red leaves is significantly lower than that of green leaves, implying an utmost effort of red leaves to increase light capture and use efficiency because of their low leaf-chlorophyll concentration. The higher petiole length of green leaves compared with that of red leaves indicates that adult green leaves may have large fractional biomass allocation to support the lamina structures in capturing light with maximum efficiency and obtaining a high growth rate. The high range of the phenotypic plasticity of leaf size, leaf thickness, single-leaf wet and dry weights, and leaf moisture of green leaves may be beneficial in achieving efficient control of water loss and nutrient deprivation. The high range of phenotypic plasticity of leaf chlorophyll concentration of red leaves may be advantageous in increasing resource (especially light) capture and use efficiency because this leaf type is juvenile in the growth stage and has low leaf-chlorophyll concentration.     

Three-dimensional lamina architecture alters light-harvesting efficiency in Fagus: a leaf-scale analysis

Modification of foliage exposition and morphology by seasonal average integrated quantum flux density (Qint) was investigated in the canopies of the shade-tolerant late-successional deciduous tree species Fagus orientalis Lipsky and Fagus sylvatica L. Because the leaves were not entirely flat anywhere in the canopy, the leaf lamina was considered to be three-dimensional and characterized by the cross-sectional angle between the leaf halves (theta). Both branch and lamina inclination angles with respect to the horizontal scaled positively with irradiance in the canopy, allowing light to penetrate to deeper canopy horizons. Lamina cross-sectional angle varied from 170 degrees in the most shaded leaves to 90-100 degrees in leaves in the top of the canopy. Thus, the degree of leaf rolling increased with increasing Qint, further reducing the light-interception efficiency of the upper-canopy leaves. Simulations of the dependence of foliage light-interception efficiency on theta demonstrated that decreases in theta primarily reduce the interception efficiency of direct irradiance, but that diffuse irradiance was equally efficiently intercepted over the entire range of theta values in our study. Despite strong alteration in foliage light-harvesting capacity within the canopy and greater transmittance of the upper crown compared with the lower canopy, mean incident irradiances varied more than 20-fold within the canopy, indicating inherent limitations in light partitioning within the canopy. This extensive canopy light gradient was paralleled by plastic changes in foliar structure and chemistry. Leaf dry mass per unit area varied 3-4-fold between the canopy top and bottom, providing an important means of scaling foliage nitrogen contents and photosynthetic capacity per unit area with Qint. Although leaf structure versus light relationships were qualitatively similar in all cases, there were important tree-to-tree and species-to-species variations, as well as evidence of differences in investments in structural compounds within the leaf lamina, possibly in response to contrasting leaf water availability in different trees.     

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

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

Structural variation in current-year shoots of broad-leaved evergreen tree saplings under forest canopies in warm temperate Japan

Stem length and leaf area of current-year shoots were measured in saplings of eight broad-leaved evergreen tree species growing under a forest canopy. Stem length varied over a range of one to two orders of magnitude within each species. In all species, both the number of leaves and the mean stem length between successive leaves were greater in longer shoots. Mean leaf size and stem length were not correlated in six of eight species, and only weakly positively correlated in the other two species. Thus, total leaf area per stem increased with stem length, but not in direct proportion: leaf area per stem length was smaller in shoots with long stems and larger in shoots with short stems. I conclude that the within-species variation in the leaf-stem balance of current-year shoots is related to variation in shoot functional roles, as has been observed for long and short shoots in many deciduous tree species: shoots with long stems are extension oriented and contribute to the framework of the crown, whereas shoots with short stems serve mainly for leaf display. Among species, large differences were found in the leaf area per stem length ratio. In the species with larger leaf area per stem length ratios, leaves had narrower blades or longer petioles, or both, resulting in a reduction of mutual shading among the leaves on the shoot.     

Photosynthesis patterns during the establishment year within two Populus clones with contrasting morphology and phenology

Diurnal and seasonal photosynthesis patterns were studied in poplar clones Populus tristis Fisch. x P. balsamfera L. cv. Tristis #1 (NC 5260) and Populus x euramericana (Dode) Guiner cv. Eugenei (NC 5326, Carolina poplar) during their first season in the field in a short rotation, intensive culture plantation. Photosynthetic rates were low in immature leaves; increased basipetally on the shoot and peaked in leaves that had recently reached full expansion; and thereafter declined in lower-crown leaves in both clones. Photosynthesis was associated with leaf age and stomatal conductance in immature leaves; adaxial photosynthetic photon flux density (PPFD) and leaf temperature in recently mature leaves; and leaf age and adaxial PPFD in lower-crown leaves. Diurnal photosynthesis patterns within trees were highly variable due to differential light interception among leaves. Results of clonal comparisons of photosynthetic rates were dependent on which leaves were pooled for comparison and how photosynthesis was expressed. Compared to Eugenei, Tristis produced smaller leaves which had higher unit-area photosynthesis rates. The more indeterminate Eugenei outgrew Tristis principally because it more fully utilized the growing season for leaf area production. Photosynthetic production integrated over the growing season was closely related to dry matter production in both clones.     

Leaf growth characteristics of fast-growing poplar hybrids Populus trichocarpa x P. deltoides

Fast-growing hybrid poplar trees (Populus trichocarpa Torr. & A. Gray x P. deltoides Bartr. ex Marsh.) were compared with slower-growing parental types in both field and laboratory experiments to determine physiological components of leaf growth that could be closely related to biomass production. Stem volume was correlated with individual leaf area (r = 0.81) and leaf growth rate (r = 0.82). Hybrids had a greater total leaf area, not because they produced more leaves, but because they had larger leaves than either parental type. The greater leaf size of the hybrids may be explained by inheritance of larger cell number from P. deltoides and larger cell size from P. trichocarpa. Rates of enlargement of isolated leaf discs in liquid culture were approximately 50% of those observed in intact leaves of field-grown plants.     

Quantitative Estimation of Genetic Variability in Morphological Characteristics of Eastern Cottonwood (Populus deltoides Bartr.) Clones

The study was carried out to analyze the genetic variability for different growth parameters in poplar clones at the age of 2 and 3 yr in the nursery. Forty-nine exotic and indigenous clones of poplar were evaluated for eight morphological traits. Clones were planted in randomized block design (RBD) with three replications with four clones in a block in each treatment. Observations for different characters were recorded on six selected competitive clones per genotype. Results showed a high interclonal variability for most parameters. Statistically significant differences among clones indicated that the majority of study characters are controlled by genetic factors, specific to each clone. Highly significant genotypic difference supported by wide range of variation of mean and range values were observed for the characters under study. Significant and positive correlation was observed between diameter at breast height (DBH) and plant height. Leaf lamina length showed positive and significant correlation with petiole length, total leaf length, leaf width, and L/B ratio. Total leaf length showed positive and significant correlation with leaf width and L/B ratio. High estimates of heritability (in a broad sense) were observed for almost all characters in the study. High genetic advance expressed as percent of mean was recorded for petiole length (60.90), followed by plant height (60.78) and collar diameter (44.19) at 2 yr. At the age of 3 yr, genetic advance was found maximum for petiole length (60.05), followed by collar diameter (47.62) and plant height (45.29). The efforts for selecting new clones and their field-testing must continue under a long-term improvement plan so that the best clones can be recommended for plantations and hybridization programs.     

Effects of Cutting Density on Growth, Yield and Quality of Poplar Clone Seedlings

In order to identify the optimum cutting density for producing the highest number of plantable seedlings of poplar clones, a split-plot randomized block design was used to establish four cutting densities in plots. Based on data on the survival, leaf area, seedling height, caliper, and biomass of 1-year-old seedlings of clones Nanlin-95, Nanlin-895, Nanlin-1388 and NL-80351, the growth characteristics and seedling quality under four cutting densities were analyzed. Results indicated that the leaf area, stem and leaf biomass, and caliper of seedlings of all four poplar clones increased with the decrease in cutting density. Leaf area index reached its highest level at the spacing of 40 cm×40 cm, while the aboveground biomass of the seedling on an area basis increased as the cutting density increased. Seedling quality at low cutting density was higher than that at closer cutting density. The quantity of first-grade seedlings (grade I) for clones Nanlin-95 and Nanlin-895 was achieved at the spacing of 40 cm×50 cm; for NL-1388 and NL-80351, it was 50 cm×50 cm. According to the seedling quality and the number of plantable seedlings produced, the suggested cutting density for these four poplar clones was 50,000 stems/hm².     

Botanical determinants of foliage clumping and light interception in two-year-old coppice poplar canopies: assessment from 3-D plant mockups

Botanical parameters (e.g., shoot and branch inclination, petiole length, leaf phyllotaxy, size and shape) that influence light interception and foliage clumping in dense two-year-old monoclonal poplar (Populus spp.) coppice crops were analysed with a three-dimensional simulation model. Crop LAI varied from 1 to 2 for clone Ghoy and from 2.5 to 7.4 for clone Trichobel from May to September. Canopies were strongly clumped, with a clumping index (μ) about 0.5. Canopy light transmittance (τ) varied from 0.59 in May to 0.41 in September for clone Ghoy and from 0.42 to 0.08 for clone Trichobel, and was strongly associated with LAI. The overall effect of a simulated shift in botanical parameters was relatively small and resulted in limited changes in μ and τ by ± 0.05 and ±0.1, respectively. Petiole length had the most notable effect on μ and τ, while the other parameters were less effective. However, biomass cost analyses showed that actual petiole length optimised the efficiency of biomass investment into light capture.     

Photosynthesis-nitrogen relationships: interpretation of different patterns between Pseudotsuga menziesii and Populus x euroamericana in a mini-stand experiment

We compared photosynthesis-nitrogen relationships of one broad-leaved (poplar; Populus x euroamericana (Dole) Guinier) and one conifer (Douglas-fir; Pseudotsuga menziesii (Mirb.) Franco) species. Plants were grown in large pots to allow free root development and were kept well watered. We determined effects of low, intermediate and high nitrogen supply rates on area-based leaf nitrogen (Na) and chlorophyll concentrations, leaf mass per area (LMA), light-saturated photosynthesis (Amax), maximum carboxylation (Vcmax) and electron transport rate (Jmax), photosynthetic nitrogen-use efficiency (PNUE), and proportions of leaf N in active Rubisco (PR), bioenergetic pools (PB) and the light-harvesting complex (PLH). Nitrogen supply significantly affected leaf Na. Leaf mass per area did not differ between species and was unaffected by the N treatments. In both species, there was a positive correlation between leaf Na and chlorophyll concentration, and between leaf Na and the photosynthetic parameters Amax, Jmax and Vcmax. At comparable leaf Na, however, poplar showed twofold higher PNUE and a threefold steeper slope of the Amax- nitrogen relationship than Douglas-fir. Leaf Na was negatively correlated with PNUE in Douglas-fir but not in poplar. Leaf Na was also negatively correlated with PR, PB and PLH in Douglas-fir, whereas in poplar, a negative correlation was found only for PLH. Parameter PR was significantly higher in poplar than in Douglas-fir. The ratio of CO2 concentration in the intercellular space to that in ambient air was higher in poplar than in Douglas-fir. Overall, our data suggest that differences in the photosynthesis-nitrogen relationship and PNUE between Douglas-fir and poplar primarily reflect a different investment of N to active Rubisco, and possibly a different constraint to CO2 diffusion.     

Morphological response of Vitex negundo var. heterophylla and Ziziphus jujuba var. spinosa to the combined impact of drought and shade

Light and soil moisture availability are two important abiotic factors influencing plant growth in an agroforestry system. Different soil moisture and light treatments were applied to examine the combined impact of drought and shade on the morphological plasticity of Vitex negundo var. heterophylla (Chinese chastetree) and Ziziphus jujuba var. spinosa (Spine jujube). We found that the interaction of light and soil moisture was orthogonal in the two species. V. negundo captured irradiance efficiently with relatively long petiole and petiolule, while Z. jujuba maintained higher branches to absorb light than V. negundo. Compared to the seedlings under full sunlight, the palmately compound leaves of V. negundo under low light showed larger specific leaf area (SLA), lower ratio of leaflet length to width and higher leaf mass ratio (leaf biomass to total biomass); in contrast, the simple leaves of Z. jujuba under low light showed larger SLA and ratio of leaf length to petiole length. In both species, drought reduced the branch number, mean internode length of stem, and increased root mass ratio (root biomass to total biomass), but leaf morphology showed little variation to the decreased soil moisture. Between the two species, V. negundo is more shade-tolerant, while Z. jujuba is more drought-tolerant. In an alley cropping system, the spacing can be smaller in the V. negundo-crop system than the Z. jujuba-crop system when soil moisture is adequate. However, the V. negundo-crop system should be managed with more caution under the soil moisture limited conditions.     

Planting spacing affects canopy structure,biomass production and stem roundness in poplar plantations

To improve the productivity and wood quality of poplar plantations, effects of four planting spacing on canopy characteristics, biomass production and stem roundness in poplar plantations were evaluated over 8 years. Planting spacing influenced canopy characteristics of the plantations, and further affected the understory vegetation and plantation productivity. Understory vegetation biomass and Shannon-Wiener index were negatively correlated with leaf area index, but both diversity indexes and aboveground biomass of understory vegetation were higher in stands with a wider spacing. Tree diameter growth increased with increasing planting spacing, while the increment in plantations of square configurations (5?×?5 m and 6?×?6 m) was higher than those with rectangular configurations (3?×?8 m, 4.5?×?8 m). The highest poplar biomass production was achieved in the plantation with 5?×?5 m spacing at age 8. Moreover, poplar trees showed a tendency with better stem roundness in a square configuration. The results suggest that planting spacing not only affect canopy characteristics, understory vegetation and tree growth but also wood quality, and square configurations (5?×?5 m and 6?×?6 m) could be a better option for poplar plywood timber production at similar sites.     

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

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

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.     

Comparison of photosynthetic induction and transient limitations during the induction phase in young and mature leaves from three poplar clones

We tested the hypothesis that leaf age affects photosynthetic induction, because conductance to CO2 diffusion usually decreases with increasing leaf age. Photosynthetic inductions, primarily determined by the light modulation of Rubisco activity and stomatal opening, were investigated in both young and mature leaves, as defined by leaf plastochron index (LPI), from three poplar clones: Populus alba L., P. nigra L. and P. x euramericana (Dode) Guinier. In all clones, maximum assimilation rates (A max), maximum stomatal conductance (G Smax) and dark respiration rates (RD) were higher in young leaves (LPI = 3-5) than in mature leaves (LPI = 10-14), and A max decreased from P. alba via P. x euramericana to P. nigra. The clones with high photosynthetic capacity had low induction states 60 s after leaf illumination (IS60; indicating a slow initial induction phase), and required less time to reach 90% photosynthetic induction (T90). In contrast, the clone with the lowest photosynthetic capacity (P. nigra) exhibited high IS60 (high initial induction state) but a long induction time (high T90). A comparison of mature leaves with young leaves revealed significantly (P < 0.01) lower IS60 values in mature leaves of P. nigra only, and significantly higher T90 values in mature leaves of P. alba only. In all clones, young leaves exhibited a lower percentage of maximum transient stomatal limitation during photosynthetic induction (4-9%) compared with mature leaves (16-30%). Transient biochemical limitation, assessed on the basis of the time constants of Rubisco activation (tau), was significantly higher in mature leaves than in young leaves of P. alba; whereas there were no significant differences in tau between young and mature leaves of the other poplar clones. Thus, our hypothesis that leaf age affects photosynthetic induction was confirmed at the level of transient stomatal limitation, which was significantly higher in mature leaves than in young leaves in all clones. For the induction parameters IS60, T90 and tau, photosynthetic induction was more clone-specific and was dependent on leaf age only in some cases, an observation that may apply to other tree species.     

The morphology and development of Lotus uliginosus and Trifolium subterraneum under Pinus radiata canopy in southern Chile

A split-plot experiment was conducted in southern Chile to study the effects of tree cover and level of fertilizer application at establishment on the morphology and development of the legumes Lotus uliginosus and Trifolium subterraneum, when sown in conjunction with Festuca arundinacea as the companion grass. Trees were 13-year-old Pinus radiata, spaced at 4 × 12.5 m intervals and with a mean canopy diameter of 4.8 m. At 150 days after sowing the total area of Lotus leaves and length of Lotus stems/plant were not affected by tree cover. Individual plants had fewer stems but they were longer. For Trifolium, despite large increases in individual petiole length and leaf area, total petiole length and leaf area/plant were reduced by tree cover because of a reduced number of petioles/plant. At 275 days after sowing, the number and weight of Lotus components/ha was not affected by tree cover, although companion grass development was reduced. By 640 days the number and weight of Lotus components/ha was reduced but only to the same degree as the companion grass. Trifolium plant development, and in particular seed yield, was affected by tree cover at 275 and 640 days and to a greater extent than Lotus or the companion grass. A high level of fertilizer application at establishment induced a greater development of both legumes at 275 days, but by 640 days the effects were much reduced. It is concluded that both Lotus uliginosus and Trifolium subterraneum respond to tree cover by increasing stem length and leaf area, but that in situations with low soil fertility, the development of Lotus is less affected by tree cover than Trifolium. This revised version was published online in June 2006 with corrections to the Cover Date.