Ontogenetic changes in stomatal and biochemical limitations to photosynthesis of two co-occurring Mediterranean oaks differing in leaf life span

A quantitative analysis was applied to the stomatal and biochemical limitations to light-saturated net photosynthesis under optimal field conditions in mature trees and seedlings of the co-occurring evergreen oak, Quercus ilex L., and the deciduous oak, Q. faginea Lam. Stomatal limitation to photosynthesis, maximal Rubisco activity and electron transport rate were determined from assimilation versus intercellular leaf carbon dioxide concentration response curves of leaves that were subsequently analyzed for nitrogen (N) concentration, mass per unit area, thickness and percent internal air space. In both species, seedlings had a lower leaf mass per unit area, thickness and leaf N concentration than mature trees. The root system of seedlings during their third year after planting was dominated by a taproot. A lower leaf N concentration of seedlings was associated with lower maximal Rubisco activity and electron transport rate and with assimilation rates similar to or lower than those of mature trees, despite the higher stomatal conductances and potential photosynthetic nitrogen-use efficiencies of seedlings. Consequently, stomatal limitation to photosynthesis increased with tree age in both species. In both seedlings and mature trees, a lower assimilation rate in Q. ilex than in Q. faginea was associated with lower stomatal conductance, N allocation to photosynthetic functions, maximal Rubisco activity and electron transport rate, and potential photosynthetic nitrogen-use efficiency but greater leaf thickness and leaf mass per unit area. Tree-age-related changes differed quantitatively between species, and the characteristics of the two species were more similar in seedlings than in mature trees. Despite higher stomatal conductances, seedlings are more N limited than adult trees, which contributes to lower biochemical efficiency.     

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.     

Gas exchange, leaf structure and nitrogen in contrasting successional tree species growing in open and understory sites during a drought

Seasonal ecophysiology, leaf structure and nitrogen were measured in saplings of early (Populus grandidentata Michx. and Prunus serotina J.F. Ehrh.), middle (Fraxinus americana L. and Carya tomentosa Nutt.) and late (Acer rubrum L. and Cornus florida L.) successional tree species during severe drought on adjacent open and understory sites in central Pennsylvania, USA. Area-based net photosynthesis (A) and leaf conductance to water vapor diffusion (g(wv)) varied by site and species and were highest in open growing plants and early successional species at both the open and understory sites. In response to the period of maximum drought, both sunfleck and sun leaves of the early successional species exhibited smaller decreases in A than leaves of the other species. Shaded understory leaves of all species were more susceptible to drought than sun leaves and had negative midday A values during the middle and later growing season. Shaded understory leaves also displayed a reduced photosynthetic light response during the peak drought period. Sun leaves were thicker and had a greater mass per area (LMA) and nitrogen (N) content than shaded leaves, and early and middle successional species had higher N contents and concentrations than late successional species. In both sunfleck and sun leaves, seasonal A was positively related to predawn leaf Psi, g(wv), LMA and N, and was negatively related to vapor pressure deficit, midday leaf Psi and internal CO(2). Although a significant amount of plasticity occurred in all species for most gas exchange and leaf structural parameters, middle successional species exhibited the largest degree of phenotypic plasticity between open and understory plants.     

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.     

Leaf optical properties in Venezuelan cloud forest trees

Leaf optical properties and related leaf characteristics were compared for thirteen cloud forest tree species differing in successional status. Sun leaves were sampled for the eight pioneer species and sun and shade leaves were sampled for the five climax species. Sun leaves had a slightly higher absorptance than shade leaves, although differences were small. Sun leaves had a higher leaf mass per unit area (LMA) and a lower chlorophyll concentration per unit leaf mass, resulting in similar chlorophyll concentrations per unit leaf area and hence similar light harvesting capacities as shade leaves. However, shade leaves realized a higher efficiency of absorptance per unit leaf biomass than sun leaves. There were few differences in leaf characteristics of sun leaves between the climax and pioneer species. Absorptance values of cloud forest species were comparable with values reported for rain forest and more seasonal forest species. Intraspecific variation in leaf absorptance was largely the result of variation in LMA, whereas interspecific variation in leaf absorptance was largely a result of variation in chlorophyll concentration per unit leaf area.     

Leaf morphological and physiological adjustments to the spectrally selective shade imposed by anthocyanins in Prunus cerasifera

Prunus domestica L. has green leaves, whereas Prunus cerasifera Ehrh. var. atropurpurea has red leaves due to the presence of mesophyll anthocyanins. We compared morphological and photosynthetic characteristics of leaves of these species, which were sampled from shoots grafted in pairs on P. domestica rootstocks, each pair comprising one shoot of each species. Two hypotheses were tested: (1) anthocyanins protect red leaves against photoinhibition; and (2) red leaves display shade characteristics because of light attenuation by anthocyanins. Parameters were measured seasonally, during a period of increasing water stress, which caused a similar drop in shoot water potential in each species. As judged by predawn measurements of maximum PSII yield, chronic photoinhibition did not develop in either species and, despite the anthocyanic screen, the red leaves of P. cerasifera displayed lower light-adapted PSII yields and higher non-photochemical quenching than the green leaves of P. domestica. Thus, it appears that, in this system, anthocyanins afford little photoprotection. As predicted by the shade acclimation hypothesis, red leaves were thinner and had a lower stomatal frequency, area- based CO2 assimilation rate, apparent carboxylation efficiency and chlorophyll a:b ratio than green leaves. However, red leaves were similar to green leaves in conductivity to water vapor diffusion, dry-mass-based chlorophyll concentrations and carotenoid:chlorophyll ratios. The data for red leaves indicate adaptations to a green-depleted, red-enriched shade, rather than a neutral or canopy-like shade. Thus, green light attenuation by anthocyanins may impose a limitation on leaf thickness. Moreover, the selective depletion of light at wavelengths that are preferentially absorbed by PSII and chlorophyll b may lead to adjustments in chlorophyll pigment ratios to compensate for the uneven spectral distribution of internal light. The apparent photosynthetic cost associated with lost photons and reduced leaf thickness, and the absence of a photoprotective advantage, suggest that there are other, yet to be identified, benefits for permanently anthocyanic leaves of P. cerasifera.     

The ecological and functional correlates of nocturnal transpiration

Contrary to the conventional theory of optimal stomatal control, there is substantial transpiration at night in many tree species, but the functional significance of this phenomenon remains uncertain. To investigate the possible roles of nocturnal transpiration, we compared and contrasted the correlations of both nocturnal and diurnal sap flow with a range of traits in 21 temperate deciduous tree species. These traits included soil water affinity, shade tolerance, cold hardiness, nitrogen concentration of tissues, minimum transpiration rate of excised leaves, growth rate, photosynthetic capacity, stomatal length and density, and the water potential and relative water content of leaves at the wilting point. Nocturnal sap flow was higher in species with higher leaf nitrogen concentrations, higher rates of extension growth and lower shade tolerances. Diurnal sap flow was higher in species with higher leaf nitrogen concentrations and photosynthetic capacities on a leaf area basis. Because leaf metabolism and dark respiration, in particular, are strongly related to leaf nitrogen concentration, our findings suggest that nocturnal transpiration functions to sustain carbohydrate export and other processes driven by dark respiration, and that this function is most important in fast- growing shade-intolerant tree species.     

Modeling photosynthesis in olive leaves under drought conditions

We quantified parameters for a model of leaf-level photosynthesis for olive, and tested the model against an independent dataset. Specific temperature-dependence parameters of the model for olive leaves were measured, as well as the relationship of the model parameters with area-based leaf nitrogen (N) content. The effect of soil water deficit on leaf photosynthesis was examined by applying two irrigation treatments to 29-year-old trees growing in a plantation: drip irrigation sufficient to meet the crop water requirements (I) and dry-farming (D). In both treatments, leaves had a higher photosynthetic capacity in April than in August. In August, photosynthetic capacity was lower in D trees than in I trees. Leaf photosynthetic capacity was linearly and positively related to leaf N content on an area basis (N(a)) and to leaf mass per unit area (LMA), and the regression slope varied with irrigation treatment. The seasonal reduction in N(a) was used in the model to predict photosynthesis under drought conditions. Olive leaves showed a clear limitation of photosynthesis by triose phosphate utilization (TPU) even at 40 degrees C, and the data suggest that olive invests fewer resources in TPU than other species. The seasonal decrease in photosynthetic capacity moderated the stomatal limitation to carbon dioxide (CO(2)) fixation as soil water deficit increased. Further, it enabled leaves to operate close to the transition point between photosynthetic limitation due to RuBP carboxylation capacity and that due to RuBP regeneration capacity, and resulted in a near constant value of internal CO(2) concentration from April to August. Under well watered conditions, N-use efficiency of the olive leaves was enhanced at the expense of reduced water-use efficiency.     

Changes in photosynthesis and water status of developing leaves of Brachystegia spiciformis Benth

Changes in net carbon assimilation and water status were studied during leaf development in the deciduous, tropical species Brachystegia spiciformis Benth. In this upland savanna African tree, bud-burst and leaf development occur approximately two months before the rainy season. The newly formed leaves synthesize anthocyanin until the fully expanded leaves of the whole canopy are red. This foliage is referred to as "spring flush" foliage. Subsequently, the anthocyanins are metabolized and the pre-rain leaves become green. Carbon dioxide assimilation exhibited a bimodal diurnal pattern and was similar for pre-rain green leaves and fully expanded flushing leaves, although pre-rain green leaves showed a net uptake of carbon throughout the daylight period, whereas flushing leaves exhibited only brief periods of net photosynthesis in the morning and early afternoon. Measurements of leaf water potential and relative water content showed a diurnal pattern with considerable variation throughout the day. Leaf water potential and relative water content values decreased soon after sunrise reaching a minimum at a time corresponding to the afternoon peak in CO(2) assimilation. Stomatal conductance was closely related to transpiration rate in both flushing and pre-rain green leaves, although flushing leaves had lower stomatal conductances than pre-rain green leaves. Pre-rain green leaves exhibited a compensation irradiance of approximately 180 micro mol m(-2) s(-1), whereas flushing leaves had positive net photosynthesis only at PPFDs greater than 300 micro mol m(-2) s(-1). Rate of photosynthesis (expressed per leaf area or chlorophyll unit) increased as anthocyanin concentration decreased, although the photosynthetic rate continued to increase long after the leaf anthocyanins had been degraded to low, visually undetectable amounts. Post-rain green leaves had chlorophyll concentrations, transpiration rates and stomatal conductances similar to those of pre-rain green leaves; however, photosynthetic rates in post-rain leaves were more than three times higher. Thus, during the early stages of the spring flush, carbon asimilation rates of the flushing leaves were inversely related to leaf anthocyanin concentrations. In pre-rain green leaves, photosynthesis was limited by other non-stomatal factors.     

Photosynthetic acclimation of overstory Populus tremuloides and understory Acer saccharum to elevated atmospheric CO2 concentration: interactions with shade and soil nitrogen

We exposed Populus tremuloides Michx. and Acer saccharum Marsh. to a factorial combination of ambient and elevated atmospheric CO2 concentrations ([CO2]) and high-nitrogen (N) and low-N soil treatments in open-top chambers for 3 years. Our objective was to compare photosynthetic acclimation to elevated [CO2] between species of contrasting shade tolerance, and to determine if soil N or shading modify the acclimation response. Sun and shade leaf responses to elevated [CO2] and soil N were compared between upper and lower canopy leaves of P. tremuloides and between A. saccharum seedlings grown with and without shading by P. tremuloides. Both species had higher leaf N concentrations and photosynthetic rates in high-N soil than in low-N soil, and these characteristics were higher for P. tremuloides than for A. saccharum. Electron transport capacity (Jmax) and carboxylation capacity (Vcmax) generally decreased with atmospheric CO2 enrichment in all 3 years of the experiment, but there was no evidence that elevated [CO2] altered the relationship between them. On a leaf area basis, both Jmax and Vcmax acclimated to elevated [CO2] more strongly in shade leaves than in sun leaves of P. tremuloides. However, the apparent [CO2] x shade interaction was largely driven by differences in specific leaf area (m2 g-1) between sun and shade leaves. In A. saccharum, photosynthesis acclimated more strongly to elevated [CO2] in sun leaves than in shade leaves on both leaf area and mass bases. We conclude that trees rooted freely in the ground can exhibit photosynthetic acclimation to elevated [CO2], and the response may be modified by light environment. The hypothesis that photosynthesis acclimates more completely to elevated [CO2] in shade-tolerant species than in shade-intolerant species was not supported.     

3种楸树1年生苗木叶片光合速率和光合功能的垂直梯度变化

对楸树、灰楸、滇楸3个树种1年生嫁接苗的净光合速率、叶面积和比叶重的垂直变化进行了测定,以探讨光合速率与苗木生长的关系。结果表明,楸树、灰楸、滇楸的净光合速率、叶面积和比叶重的垂直变化趋势基本相似,但树种之间的波动幅度上存在着差异,灰楸的总体光合速率较低,滇楸前期叶面积增加速度明显大于楸树和灰楸,而后期下降趋势明显低于楸树和灰楸,且下部叶片能维持较高的光合速率,最末叶序的比叶重都比刚展叶时的比叶重高。苗高生长量与除比叶重外的4个指标呈极显著正相关,光合指标与5个指标都呈极显著正相关。     

Seasonal patterns of leaf gas exchange and water relations in dry rain forest trees of contrasting leaf phenology

Diurnal and seasonal patterns of leaf gas exchange and water relations were examined in tree species of contrasting leaf phenology growing in a seasonally dry tropical rain forest in north-eastern Australia. Two drought-deciduous species, Brachychiton australis (Schott and Endl.) A. Terracc. and Cochlospermum gillivraei Benth., and two evergreen species, Alphitonia excelsa (Fenzal) Benth. and Austromyrtus bidwillii (Benth.) Burret. were studied. The deciduous species had higher specific leaf areas and maximum photosynthetic rates per leaf dry mass in the wet season than the evergreens. During the transition from wet season to dry season, total canopy area was reduced by 70-90% in the deciduous species and stomatal conductance (g(s)) and assimilation rate (A) were markedly lower in the remaining leaves. Deciduous species maintained daytime leaf water potentials (Psi(L)) at close to or above wet season values by a combination of stomatal regulation and reduction in leaf area. Thus, the timing of leaf drop in deciduous species was not associated with large negative values of daytime Psi(L) (greater than -1.6 MPa) or predawn Psi(L) (greater than -1.0 MPa). The deciduous species appeared sensitive to small perturbations in soil and leaf water status that signalled the onset of drought. The evergreen species were less sensitive to the onset of drought and g(s) values were not significantly lower during the transitional period. In the dry season, the evergreen species maintained their canopies despite increasing water-stress; however, unlike Eucalyptus species from northern Australian savannas, A and g(s) were significantly lower than wet season values.     

Leaf gas exchange in the frankincense tree (Boswellia papyrifera) of African dry woodlands

A conceptual model was tested for explaining environmental and physiological effects on leaf gas exchange in the deciduous dry tropical woodland tree Boswellia papyrifera (Del.) Hochst. For this species we aimed at (i) understanding diurnal patterns in leaf gas exchange, (ii) exploring cause-effect relationships among external environment, internal physiology and leaf gas exchange, and (iii) exploring site differences in leaf gas exchange in response to environmental variables. Diurnal courses in gas exchange, underlying physiological traits and environmental variables were measured for 90 trees on consecutive days at two contrasting areas, one at high and the other at low altitude. Assimilation was highest in the morning and slightly decreased during the day. In contrast, transpiration increased from early morning to midday, mainly in response to an increasing vapor pressure deficit (VPD) and gradual stomatal closure. The leaf water potential varied relatively little and did not influence gas exchange during the measurement period. Our results suggest that the same cause-effect relationships function at contrasting areas. However, leaves at the higher altitude had higher photosynthetic capacity, reflecting acclimation to higher light levels. Trees at both areas nevertheless achieved similar leaf assimilation rates since assimilation was down-regulated by stomatal closure due to the higher VPD at the higher altitude, while it became more light limited at the lower altitude. Gas exchange was thus limited by a high VPD or low light levels during the wet season, despite the ability of the species to acclimate to different conditions.     

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.     

Photosynthesis and chlorophylla fluorescence of two poplars under water stress

In order to estimate drought tolerance in two species ofPopulus, Populus alba var. pyramidalis Bunge andPopulus nigra L. var. thevestina (Dode), widely planted at the southern margin of the Taklimakan Desert, responses of net photosynthesis and chlorophylla fluorescence to irradiance and water stress were examined under laboratory conditions. Results showed thatP. alba exhibited stronger drought tolerance thanP. nigra. A linear relationship between net photosynthetic rates (A _n) and electron transport rates (ETR) was found in both poplars under different irradiance and leaf water potentials. Net photosynthetic rates (A _n) in the two poplars significantly correlated linearly with the photochemical efficiency of the saturation light-adapted leaves throughout the range of leaf water potentials, suggesting that the leaf photochemical efficiency in saturation light-adapted leaves can be used to estimate leaf photosynthetic capacity and leaf water conditions in the two poplars within a magnitude of air temperature between 20 and 30°C. This research was supported by the Japan Society for the Promotion of Science (CAS-9601), the Chinese Academy of Sciences and partly supported by the National Natural Foundation of China (39870154).     

Influence of overstory density on ecophysiology of red oak (Quercus rubra) and sugar maple (Acer saccharum) seedlings in central Ontario shelterwoods

A field experiment was established in a second-growth hardwood forest dominated by red oak (Quercus rubra L.) to examine the effects of shelterwood overstory density on leaf gas exchange and seedling water status of planted red oak, naturally regenerated red oak and sugar maple (Acer saccharum Marsh.) seedlings during the first growing season following harvest. Canopy cover of uncut control stands and moderate and light shelterwoods averaged 97, 80 and 49%, respectively. Understory light and vapor pressure deficit (VPD) strongly influenced gas exchange responses to overstory reduction. Increased irradiance beneath the shelterwoods significantly increased net photosynthesis (P(n)) and leaf conductance to water vapor (G(wv)) of red oak and maple seedlings; however, P(n) and G(wv) of planted and naturally regenerated red oak seedlings were two to three times higher than those of sugar maple seedlings in both partial harvest treatments, due in large part to decreased stomatal limitation of gas exchange in red oak as a result of increased VPD in the shelterwoods. In both species, seedling water status was higher in the partial harvest treatments, as reflected by the higher predawn leaf water potential and seedling water-use efficiency in seedlings in shelterwoods than in uncut stands. Within a treatment, planted and natural red oak seedlings exhibited similar leaf gas exchange rates and water status, indicating little adverse physiological effect of transplanting. We conclude that the use of shelterwoods favors photosynthetic potential of red oak over sugar maple, and should improve red oak regeneration in Ontario.     

Moderate shade can increase net gas exchange and reduce photoinhibition in citrus leaves

Daily variations in net gas exchange, chlorophyll a fluorescence and water relations of mature, sun-acclimated grapefruit (Citrus paradisi Macfady.) and orange (Citrus sinensis L. Osbeck) leaves were determined in tree canopies either shaded with 50% shade screens or left unshaded (sunlit). Mean daily maximum photosynthetic photon flux density (PPFD) under shade varied from 500 to 700 micromol m-2 s-1 and was sufficient to achieve maximum net CO2 assimilation rates (A CO2). Responses of grapefruit and orange leaves to shading were remarkably similar. At midday, on bright clear days, the temperatures of sunlit leaves were 2-6 degrees C above air temperature and 1-4 degrees C above the temperatures of shaded leaves. Although midday depressions of stomatal conductance (gs) and A CO2 were observed in both sunlit and shaded leaves, shaded leaves had lower leaf-to-air vapor pressure differences (D) along with higher gs, A CO2 and leaf water-use efficiency than sunlit leaves. Estimated stomatal limitation to A CO2 was generally less than 25% and did not differ between shaded and sunlit leaves. Leaf intercellular CO2 partial pressure was not altered by shade treatment and did not change substantially with increasing D. Radiation and high temperature stress-induced non-stomatal limitation to A CO2 in sunlit leaves was greater than 40%. Reversible photoinhibition of photosystem II efficiency was more pronounced in sunlit than in shaded leaves. Thus, non-stomatal factors play a major role in regulating A CO2 of citrus leaves during radiation and high temperature stress.     

Response of Ulmus americana seedlings to varying nitrogen and water status. 1 Photosynthesis and growth

Well-watered American elm (Ulmus americana L.) seedlings responded to increased nitrate availability with increased leaf nitrogen (N) concentration and photosynthetic rate, larger and more numerous leaves, greater total growth and greater proportional allocation of carbon to shoot than root. Plasticity of growth and carbon allocation were greater than plasticity of N concentration and photosynthetic capacity. For a given N availability, allocation of N per unit leaf area was positively correlated with dry mass per unit leaf area (specific leaf mass), but these relationships differed with N availability. Rates of net photosynthesis and leaf conductance declined logarithmically with decreasing predawn water status. Increased water stress resulted in a greater relative decline in net photosynthesis and leaf conductance for high-N than low-N plants.     

Seasonal changes in photosynthesis of trees in the flooded forest of the Mapire River

We studied the flood tolerance of five tree species growing in the flooded forest adjacent to the Mapire river, in SW Venezuela. Mean photosynthetic rate and leaf conductance were 11 &mgr;mol m(-2) s(-1) and 700 mmol m(-2) s(-1), respectively. Xylem water potential ranged from -0.08 to -1.15 MPa. Based on leaf gas exchange as a criterion of tolerance to flooding, two response patterns were identified: (1) decreasing photosynthetic rate with increasing flooding and leaf conductance (Psidium ovatifolium Berg. ex Desc., Campsiandra laurifolia Benth., Symmeria paniculata Benth. and Acosmium nitens (Vog.) Benth); and (2) independence of photosynthesis and leaf conductance from flooding (Eschweilera tenuifolia (Berg.) Miers.). In the first response pattern, declining photosynthetic rate with flooding may be interpreted as a sign of reduced flood tolerance, whereas the second response pattern may indicate increased flood tolerance. An increase in xylem water potential with depth of water column was found for all species (with the possible exception of P. ovatifolium), indicating that flooding does not cause water stress in these trees. Submerged leaves that had been under water for between four days and four months generally had photosynthetic rates and leaf conductances similar to those of aerial leaves, indicating maintenance of photosynthetic capacity under water. Daily positive oscillations in glucan content in submerged leaves of P. ovatifolium and C. laurifolia suggest that submerged leaves do not represent a sink for photosynthates produced by aerial leaves.     

Contributions of foliage distribution and leaf functions to light interception, transpiration and photosynthetic capacities in two apple cultivars at branch and tree scales

Both the spatial distribution of leaves and leaf functions affect the light interception, transpiration and photosynthetic capacities of trees, but their relative contributions have rarely been investigated. We assessed these contributions at the branch and tree scales in two apple cultivars (Malus x domestica Borkh. 'Fuji' and 'Braeburn') with contrasting architectures, by estimating their branch and tree capacities and comparing them with outputs from a radiation absorption, transpiration and photosynthesis (RATP) functional-structural plant model (FSPM). The structures of three 8-year-old trees of each cultivar were digitized to obtain 3-D representations of foliage geometry. Within-tree foliage distribution was compared with shoot demography, number of leaves per shoot and mean individual leaf area. We estimated branch and tree light interception from silhouette to total leaf area ratios (STAR), transpiration from sap flux measurements and net photosynthetic rates by the branch bag method. Based on a set of parameters we previously established for both cultivars, the outputs of the RATP model were tested against STAR values, sap fluxes and photosynthetic measurements. The RATP model was then used to virtually switch foliage distribution or leaf functions (stomatal and photosynthetic properties), or both, between cultivars and to evaluate the effects on branch and tree light interception, transpiration and photosynthetic capacities in each cultivar. 'Fuji' trees had a higher proportion of leaf area borne on long shoots, fewer leaves per unit shoot length and a larger individual leaf area than 'Braeburn' trees. This resulted in a lower leaf area density and, consequently, a higher STAR in 'Fuji' than in 'Braeburn' at both branch and tree scales. Transpiration and photosynthetic rates were significantly higher in 'Fuji' than in 'Braeburn'. Branch heterogeneity was greater in 'Braeburn' than in 'Fuji'. An analysis of the virtual switches of foliage distribution or leaf function showed that differences in leaf spatial distribution and functions had additive effects that accounted for the lower transpiration and photosynthetic rates of branches and trees of 'Braeburn' compared with 'Fuji'. Leaf distribution had a more important role at the branch scale than at the tree scale, but the leaf function effect exceeded the leaf distribution effect at both scales. Our study demonstrated the potential of FSPM to disentangle physiological differences between cultivars through in silico scenarios.