Do the capacity and kinetics for modification of xanthophyll cycle pool size depend on growth irradiance in temperate trees?

The present study investigated the interaction of growth irradiance (Q_int) with leaf capacity for and kinetics of adjustment of the pool size of xanthophyll cycle carotenoids (sum of violaxanthin, antheraxanthin and zeaxanthin; VAZ) and photosynthetic electron transport rate (J_max) after changes in leaf light environment. Individual leaves of lower‐canopy/lower photosynthetic capacity species Tilia cordata Mill. and upper canopy/higher photosynthetic capacity species Populus tremula L. were either illuminated by additional light of 500–800 µmol m^?2 s^?1 for 12 h photoperiod or enclosed in shade bags. The extra irradiance increased the total amount of light intercepted by two‐fold for the upper and 10–15‐fold for the lower canopy leaves, whereas the shade bags transmitted 45% of incident irradiance. In control leaves, VAZ/area, VAZ/Chl and J_max were positively associated with leaf growth irradiance (Q_int). After 11 d extra illumination, VAZ/Chl increased in all cases due to a strong reduction in foliar chlorophyll, but VAZ/area increased in the upper canopy leaves of both species, and remained constant or decreased in the lower canopy leaves of T. cordata. The slope for VAZ/area changes with cumulative extra irradiance was positively associated with Q_int only in T. cordata, but not in P. tremula. Nevertheless, all leaves of P. tremula increased VAZ/area more than the most responsive leaves of T. cordata. Shading reduced VAZ content only in P. tremula, but not in T. cordata, again demonstrating that P. tremula is a more responsive species. Compatible with the hypothesis of the role of VAZ in photoprotection, the rates of photosynthetic electron transport declined less in P. tremula than in T. cordata after the extra irradiance treatment. However, foliar chlorophyll contents of the exposed leaves declined significantly more in the upper canopy of P. tremula, which is not consistent with the suggestion that the leaves with the highest VAZ content are more resistant to photoinhibition. This study demonstrates that previous leaf light environment may significantly affect the adaptation capacity of foliage to altered light environment, and also that species differences in photosynthetic capacity and acclimation potentials importantly alter this interaction.     

Light capture efficiency decreases with increasing tree age and size in the southern hemisphere gymnosperm Agathis australis

We investigated leaf and shoot architecture in relation to growth irradiance (Q_int) in young and mature trees of a New Zealand native gymnosperm Agathis australis (D. Don) Lindl. to determine tree size-dependent and age-dependent controls on light interception efficiency. A binomial 3-D turbid medium model was constructed to distinguish between differences in shoot light interception efficiency due to variations in leaf area density, angular distribution and leaf aggregation. Because of the positive effect of light on leaf dry mass per area (M_A), nitrogen content per area (N_A) increased with increasing irradiance in both young and mature trees. At a common irradiance, N_A, M_A and the components of M_A, density and thickness, were larger in mature trees, indicating a greater accumulation of photosynthetic biomass per unit area, but also a larger fraction of support biomass in older trees. In both young and mature trees, shoot inclination angle relative to horizontal, and leaf number per unit stem length decreased, and silhouette to total leaf area ratio (S_S) increased with decreasing irradiance, demonstrating more efficient light harvesting in low light. The shoots of young trees were more horizontal and less densely leafed with a larger S_S than those of mature trees, signifying greater light interception efficiency in young plants. Superior light harvesting in young trees resulted from more planar leaf arrangement and less clumped foliage. These results suggest that the age-dependent and/or size-dependent decreases in stand productivity may partly result from reduced light interception efficiency in larger mature relative to smaller and younger plants.     

Adjustment of foliage structure and function to a canopy light gradient in two co-existing deciduous trees. Variability in leaf inclination angles in relation to petiole morphology

 Foliar inclination angles, petiole morphology and dry matter partitioning between assimilative and support biomass were studied in shade-intolerant Populus tremula L. and shade-tolerant Tilia cordata Mill. along a natural light gradient across the canopy. The leaves of sub-canopy species T. cordata were on average exposed to lower irradiances, and they were also more horizontal with greater blade inclination angles (ϕ_B, defined as the angle between the leaf fall-line and the horizon; ϕ_B was positive for the leaves inclined upwards, and negative for the leaves inclined downwards) than those in P. tremula. Seasonal average daily integrated quantum flux density (Q _int, mol m^–2 day^–1) and ϕ_B were not related in T. cordata, and only a weak negative effect of Q _int on ϕ_B was detected in P. tremula. Nevertheless, when both species were pooled, there was a strong negative relationship between Q _int and ϕ_B, implying that the leaves became progressively vertical with increasing height in the canopy. Interspecific differences in foliage inclination were mainly related to petiole morphology, in particular to petiole length, rather than to contrasting biomass investment patterns between assimilative and support tissues within the leaf. It was suggested that more horizontal leaves, resulting from the species-specific structure of petioles, partly explain the superior performance of shade-tolerant T. cordata in the understory and the sub-canopy. Received: 13 November 1997 / Accepted: 6 March 1998     

Canopy gradients in leaf intercellular CO2 mole fractions revisited: interactions between leaf irradiance and water stress need consideration

Intercellular CO₂ mole fractions (C_i) are lower in the upper canopy relative to the lower canopy leaves. This canopy gradient in C_i has been associated with enhanced rates of carbon assimilation at high light, and concomitant greater draw‐downs in C_i. However, increases in irradiance in the canopy are generally also associated with decreases in leaf water availability. Thus, stress effects on photosynthesis rates (A) and stomatal conductance (G), may provide a further explanation for the observed C_i gradients. To test the hypotheses of the sources of canopy variation in C_i, and quantitatively assess the influence of within‐canopy differences in stomatal regulation on A, the seasonal and diurnal variation in G was studied in relation to seasonal average daily integrated quantum flux density (Q_int) in tall shade‐intolerant Populus tremula L. trees. Daily time‐courses of A were simulated using the photosynthesis model of Farquhar et al. (Planta 149, 78–90, 1980). Stable carbon isotope composition of a leaf carbon fraction with rapid turnover rate was used to estimate canopy gradient in C_i during the simulations. Daily maximum G (G_max) consistently increased with increasing Q_int. However, canopy differences in G_max decreased as soil water availability became limiting during the season. In water‐stressed leaves, there were strong mid‐day decreases in G that were poorly associated with vapour pressure deficits between the leaf and atmosphere, and the magnitude of the mid‐day decreases in G occasionally interacted with long‐term leaf light environment. Simulations indicated that the percentage of carbon lost due to mid‐day stomatal closure was of the order of 5–10%, and seasonal water stress increased this percentage up to 20%. The percentage of carbon lost due to stomatal closure increased with increasing Q_int. Canopy differences in light environment resulted in a gradient of daily average C_i of approximately 20 µmol mol^?1. The canopy variation in seasonal and diurnal reductions in G led to a C_i gradient of approximately 100 µmol mol^?1, and the actual canopy C_i gradient was of the same magnitude according to leaf carbon isotope composition. This study demonstrates that stress effects influence C_i more strongly than within‐canopy light gradients, and also that leaves acclimated to different irradiance and water stress conditions may regulate water use largely independent of foliar photosynthetic potentials.     

Petiole mechanics, leaf inclination, morphology, and investment in support in relation to light availability in the canopy of Liriodendron tulipifera

To determine the role of leaf mechanical properties in altering foliar inclination angles, and the nutrient and carbon costs of specific foliar angle variation patterns along the canopy, leaf structural and biomechanical characteristics, biomass partitioning into support, and foliar nitrogen and carbon concentrations were studied in the temperate deciduous species Liriodendron tulipifera L., which possesses large leaves on long petioles. We used beam theory to model leaf lamina as a uniform load, and estimated both the lamina and petiole flexural stiffness, which characterizes the resistance to bending of foliar elements at a common load and length. Petiole and lamina vertical inclination angles with respect to horizontal increased with increasing average daily integrated photon flux density (Q_int). Yet, the light effects on lamina inclination angle were primary determined by the petiole inclination angle. Although the petioles and laminas became longer, and the lamina loads increased with increasing Q_int, the flexural stiffness of both lamina and petiole increased to compensate for this, such that the lamina vertical displacement was only weakly related to Q_int. In addition, increases and decreases in the petiole inclination angle with respect to the horizontal effectively reduced the distance of lamina load from the axis of rotation, thereby reducing the bending moments and lamina inclination due to gravity. We demonstrate that large investments, up to 30% of total leaf biomass, in petiole and large veins are necessary to maintain the lamina at a specific position, but also that light has no direct effect on the fractional biomass investment in support. However, we provide evidence that apart from light availability, structural and chemical characteristics of the foliage may also be affected by water stress, magnitude of which scales positively with Q_int.     

Within-canopy variation in the rate of development of photosynthetic capacity is proportional to integrated quantum flux density in temperate deciduous trees

The present study was undertaken to test for the hypothesis that the rate of development in the capacity for photosynthetic electron transport per unit area (J_max;A), and maximum carboxylase activity of Rubisco (V_cmax;A) is proportional to average integrated daily quantum flux density (Q_int) in a mixed deciduous forest dominated by the shade‐intolerant species Populus tremula L., and the shade‐tolerant species Tilia cordata Mill. We distinguished between the age‐dependent changes in net assimilation rates due to modifications in leaf dry mass per unit area (M_A), foliar nitrogen content per unit dry mass (N_M), and fractional partitioning of foliar nitrogen in the proteins of photosynthetic electron transport (F_B), Rubisco (F_R) and in light‐harvesting chlorophyll‐protein complexes (V_cmax;A ∝ M_AN_MF_R; J_max;A ∝ M_AN_MF_B). In both species, increases in J_max;A and V_cmax;A during leaf development were primarily determined by nitrogen allocation to growing leaves, increases in leaf nitrogen partitioning in photosynthetic machinery, and increases in M_A. Canopy differences in the rate of development of leaf photosynthetic capacity were mainly controlled by the rate of change in M_A. There was only small within‐canopy variation in the initial rate of biomass accumulation per unit Q_int (slope of M_A versus leaf age relationship per unit Q_int), suggesting that canopy differences in the rate of development of J_max;A and V_cmax;A are directly proportional to Q_int. Nevertheless, M_A, nitrogen, J_max;A and V_cmax;A of mature leaves were not proportional to Q_int because of a finite M_A in leaves immediately after bud‐burst (light‐independent component of M_A). M_A, leaf chlorophyll contents and chlorophyll : N ratio of mature leaves were best correlated with the integrated average quantum flux density during leaf development, suggesting that foliar photosynthetic apparatus, once developed, is not affected by day‐to‐day fluctuations in Q_int. However, for the upper canopy leaves of P. tremula and for the entire canopy of T. cordata, there was a continuous decline in N contents per unit dry mass in mature non‐senescent leaves on the order of 15–20% for a change of leaf age from 40 to 120 d, possibly manifesting nitrogen reallocation to bud formation. The decline in N contents led to similar decreases in leaf photosynthetic capacity and foliar chlorophyll contents. These data demonstrate that light‐dependent variation in the rate of developmental changes in M_A determines canopy differences in photosynthetic capacity, whereas foliar photosynthetic apparatus is essentially constant in fully developed leaves.     

Shape of leaf photosynthetic electron transport versus temperature response curve is not constant along canopy light gradients in temperate deciduous trees

Responses of foliar light-saturated net assimilation rate (A_max), capacity for photosynthetic electron transport (J_max) and mitochondrial respiration rate (R_d) to long-term canopy light and temperature environment were investigated in a temperate deciduous canopy composed of Populus tremula L. in the upper (17–28 m) and of Tilia cordata Mill. in the lower canopy layer (4–17 m). Climatic measurements indicated that seasonal average daily maximum air temperature (T_max) was 5·5 °C (range 0·7–10·5 °C) higher in the top than in the bottom of the canopy, and strong positive correlations were observed between T_max and seasonal average integrated quantum flux density (Q_int), as well as between seasonal average daily mean temperature and Q_int. Because of changes in leaf dry mass and nitrogen per unit area, A_max, J_max, and R_d scaled positively with Q_int in both species at a common leaf temperature (T). According to J_max versus T response curves and dark chlorophyll fluorescence transients, photosynthetic electron transport was less heat resistant in P. tremula with optimum temperature of J_max, T_opt, of 33·5 ± 0·6 °C than in T. cordata with T_opt of 40·7 ± 0·6 °C. This difference was suggested to manifest evolutionary adaptation of photosynthetic electron transport to cooler environments in P. tremula, the range of which extends farther north than that in T. cordata. Possibly because of acclimation to long-term canopy temperature environment, T_opt was positively related to Q_int in P. tremula, foliage of which was also exposed to higher irradiances and temperatures, but not in T. cordata, in the canopy of which quantum flux densities and temperatures were lower, and gradients in the environmental factors less pronounced. Parallel to changes in T_opt, the activation energy for photosynthetic electron transport decreased with increasing Q_int in P. tremula, indicating that J_max of leaves acclimated to colder environment was more responsive to T in lower temperatures than that of high T acclimated leaves. Similar alterations in the activation energy for mitochondrial respiration rate were also observed, indicating that acclimation to temperature of mitochondrial and chloroplastic electron transport proceeds in a co-ordinated manner, and possibly involves long-term changes in membrane fluidity properties. We conclude that, because of correlations between temperature and light, the shapes of J_max versus T, and R_d versus T response curves vary within tree canopies, and this needs to be taken account in modelling whole canopy photosynthesis.     

Effects of light availability versus hydraulic constraints on stomatal responses within a crown of silver birch

Responses of leaf conductance (g_L) to variation in photosynthetic photon flux density (Q_P), leaf-to-air vapour pressure difference (VPD), bulk leaf water potential (_x), and total hydraulic conductance (G_T) were examined in silver birch (Betula pendula Roth) with respect to leaf position in the crown. To reduce limitations caused by insufficient water supply or low light availability, experiments were also performed with branchlets cut from two different canopy layers. The intact upper-canopy leaves demonstrated 1.8–2.0 times higher (P<0.001) daily maxima of g_L compared with the lower-canopy leaves growing in the shadow of upper branches. In the morning, g_L in the shade foliage was primarily constrained by low light availability, in the afternoon, by limited water supply. Leaf conductance decreased when _x fell below certain values around midday, while the sun foliage experienced greater negative water potentials than the shade foliage. Midday stomatal openness was controlled by leaf water status and temperature, rather than by transpiration rate (E) via the feedforward mechanism. Mean G_T was 1.7 times higher (P<0.001) for the upper-canopy foliage compared to that of the lower canopy. At least 34–39% of the total resistance to the water flow from soil up to the shade foliage, and 54% up to the sun foliage, resided in 30-cm distal parts of the branches. Artificial reduction of hydraulic constraints raised _x and made g_L less sensitive to changes in both atmospheric and plant factors. Improved water supply increased g_L and E in the lower-canopy foliage, but not in the upper-canopy foliage. The results support the idea that leaves in the lower canopy are hydraulically more constrained than in the upper canopy.     

A review of light interception in plant stands from leaf to canopy in different plant functional types and in species with varying shade tolerance

Changes in the efficiency of light interception and in the costs for light harvesting along the light gradients from the top of the plant canopy to the bottom are the major means by which efficient light harvesting is achieved in ecosystems. In the current review analysis, leaf, shoot and canopy level determinants of plant light harvesting, the light-driven plasticity in key traits altering light harvesting, and variations among different plant functional types and between species of different shade tolerance are analyzed. In addition, plant age- and size-dependent alterations in light harvesting efficiency are also examined. At the leaf level, the variations in light harvesting are driven by alterations in leaf chlorophyll content modifies the fraction of incident light harvested by given leaf area, and in leaf dry mass per unit area (M _A) that determines the amount of leaf area formed with certain fraction of plant biomass in the leaves. In needle-leaved species with complex foliage cross-section, the degree of foliage surface exposure also depends on the leaf total-to-projected surface area ratio. At the shoot scale, foliage inclination angle distribution and foliage spatial aggregation are the major determinants of light harvesting, while at the canopy scale, branching frequency, foliage distribution and biomass allocation to leaves (F _L) modify light harvesting significantly. F _L decreases with increasing plant size from herbs to shrubs to trees due to progressively larger support costs in plant functional types with greater stature. Among trees, F _L and stand leaf area index scale positively with foliage longevity. Plant traits altering light harvesting have a large potential to adjust to light availability. Chlorophyll per mass increases, while M _A, foliage inclination from the horizontal and degree of spatial aggregation decrease with decreasing light availability. In addition, branching frequency decreases and canopies become flatter in lower light. All these plastic modifications greatly enhance light harvesting in low light. Species with greater shade tolerance typically form a more extensive canopy by having lower M _A in deciduous species and enhanced leaf longevity in evergreens. In addition, young plants of shade tolerators commonly have less strongly aggregated foliage and flatter canopies, while in adult plants partly exposed to high light, higher shade tolerance of foliage allows the shade tolerators to maintain more leaf layers, resulting in extended crowns. Within a given plant functional type, increases in plant age and size result in increases in M _A, reductions in F _L and increases in foliage aggregation, thereby reducing plant leaf area index and the efficiency of light harvesting. Such dynamic modifications in plant light harvesting play a key role in stand development and productivity. Overall, the current review analysis demonstrates that a suite of chemical and architectural traits at various scales and their plasticity drive plant light harvesting efficiency. Enhanced light harvesting can be achieved by various combinations of traits, and these suites of traits vary during plant ontogeny.     

Changes in foliage distribution with relative irradiance and tree size: Differences between the saplings ofAcer platanoides andQuercus robur

Dependencies of foliage arrangement and structure on relative irradiance and total height (TH) were studied in saplings ofAcer platanoides andQuercus robur. The distribution of relative foliar area and dry weight (leaf area and weight in a crown layer per total tree leaf area and weight, respectively) were examined with respect to relative height (RH, height in the crown per TH) and characterized by the Weibull function. The distributions of relative area and weight were nearly identical, and the differences between them were attributable to a systematic decline in leaf dry weight per area with increasing crown depth. Foliage distribution was similarly altered by tree size in both species; RH at foliage maximum was lower and relative canopy size (RCS, length of live crown per TH) greater in taller trees. However, the distribution was more uniform inA. platanoides than inQ. robur. Apart from the size effects, relative irradiance also influenced canopy structure; RCS increased inQ. platanoides and decreased inQ. robur with increasing irradiance. As crown architecture was modified by irradiance, foliage distribution was shifted upwards with decreasing irradiance inA. platanoides, but it was independent of irradiance inQ. robur. Higher foliage maximum at lower irradiance in more shade-tolerantA. platanoides is likely to contribute towards more efficient foliar display for light interception and increase the competitive ability of this species in light-limited environments. Consequently, these differences in crown architecture and foliage distribution may partly explain the superior behavior ofA. platanoides in understory.     

Acclimation of photosynthetic capacity to irradiance in tree canopies in relation to leaf nitrogen concentration and leaf mass per unit area

The observation of acclimation in leaf photosynthetic capacity to differences in growth irradiance has been widely used as support for a hypothesis that enables a simplification of some soil‐vegetation‐atmosphere transfer (SVAT) photosynthesis models. The acclimation hypothesis requires that relative leaf nitrogen concentration declines with relative irradiance from the top of a canopy to the bottom, in 1 : 1 proportion. In combination with a light transmission model it enables a simple estimate of the vertical profile in leaf nitrogen concentration (which is assumed to determine maximum carboxylation capacity), and in combination with estimates of the fraction of absorbed radiation it also leads to simple ‘big‐leaf’ analytical solutions for canopy photosynthesis. We tested how forests deviate from this condition in five tree canopies, including four broadleaf stands, and one needle‐leaf stand: a mixed‐species tropical rain forest, oak (Quercus petraea (Matt.) Liebl), birch (Betula pendula Roth), beech (Fagus sylvatica L.) and Sitka spruce (Picea sitchensis (Bong.) Carr). Each canopy was studied when fully developed (mid‐to‐late summer for temperate stands). Irradiance (Q, µmol m^?2 s^?1) was measured for 20 d using quantum sensors placed throughout the vertical canopy profile. Measurements were made to obtain parameters from leaves adjacent to the radiation sensors: maximum carboxylation and electron transfer capacity (V_a, J_a, µmol m^?2 s^?1), day respiration (R_da, µmol m^?2 s^?1), leaf nitrogen concentration (N_m, mg g^?1) and leaf mass per unit area (L_a, g m^?2). Relative to upper‐canopy values, V_a declined linearly in 1 : 1 proportion with N_a. Relative V_a also declined linearly with relative Q, but with a significant intercept at zero irradiance (P < 0·01). This intercept was strongly related to L_a of the lowest leaves in each canopy (P < 0·01, r² = 0·98, n= 5). For each canopy, daily lnQ was also linearly related with lnV_a(P < 0·05), and the intercept was correlated with the value for photosynthetic capacity per unit nitrogen (PUN: V_a/N_a, µmol g^?1 s^?1) of the lowest leaves in each canopy (P < 0·05). V_a was linearly related with L_a and N_a(P < 0·01), but the slope of the V_a : N_a relationship varied widely among sites. Hence, whilst there was a unique V_a : N_a ratio in each stand, acclimation in V_a to Q varied predictably with L_a of the lowest leaves in each canopy. The specific leaf area, L_m(cm² g^?1), of the canopy‐bottom foliage was also found to predict carboxylation capacity (expressed on a mass basis; V_m, µmol g^?1 s^?1) at all sites (P < 0·01). These results invalidate the hypothesis of full acclimation to irradiance, but suggest that L_a and L_m of the most light‐limited leaves in a canopy are widely applicable indicators of the distribution of photosynthetic capacity with height in forests.     

Is distribution of hydraulic constraints within tree crowns reflected in photosynthetic water-use efficiency? An example of <Emphasis Type="Italic">Betula pendula</Emphasis>

Spatial and daily variation in photosynthetic water-use efficiency was examined in leaves of Betula pendula Roth with respect to distribution of hydraulic conductance within the crown, morphological properties of stomata, and water availability. Intrinsic water-use efficiency (A _n/g _s) was determined from gas-exchange measurements performed both in situ in a natural forest stand and on detached shoots under laboratory conditions. In intact foliage, sun leaves demonstrated significantly higher (P < 0.001) A _n/g _s than shade leaves, as photosynthesis in the lower canopy was chronically limited by low light availability. However, this difference reversed in the mid-day period under sufficient irradiance (I > 800 μmol m⁻² s⁻¹): A _n/g _s averaged 28.8 and 24.0 μmol mol⁻¹ (P < 0.01) for shade and sun leaves, respectively. This last finding coincided with the data obtained in laboratory conditions: under equivalent leaf water supply and light, A _n/g _s in shade foliage was greater (P < 0.001) than in sun foliage across a wide range of irradiance. Thus, shade foliage of B. pendula is characterized by inherently higher A _n/g _s than sun foliage, associated with more conservative stomatal behavior, and lower soil-to-leaf (K _T) and leaf hydraulic conductances. Under unlimited light conditions, a within-crown trade-off between A _n/g _s and K _T becomes apparent. Differences in stomatal conductance between the detached shoots from sunlit and shaded canopy layers were largely attributable to the variation in stomatal morphology; significant relationships were established with characteristics combining stomatal size and density (relative stomatal surface, stomatal pore area index). Stomatal morphology is very likely involved in long-term adjustment of photosynthetic WUE.     

Contrasting correlation networks between leaf structure,nitrogen and chlorophyll in herbaceous and woody canopies

We studied acclimation patterns in leaf dry mass per area (M_A), nitrogen (N_A) and chlorophyll (ζ_A) content per area, and chlorophyll to nitrogen ratio (ζ/N) along vertical light gradients in natural temperate mixed herbaceous canopy and deciduous tree canopy. In the deciduous tree canopy, all leaves are formed at approximately the same time, and the light gradient during the rest of the growing season reflects the differences in light availability during leaf development, whereas in the herbaceous canopy, leaf production continues during most of the growing season and major changes in light conditions occur after leaf maturation. M_A and N_A increased strongly with increasing current light availability (I_D) in the tree canopy. In the herbaceous canopy, M_A and N_A were generally unrelated to I_D. Depending on species, the correlation between chlorophyll content per leaf area (ζ_A) and I_D was positive, negative, or non-significant. Path analyses revealed two opposite effects of I_D on the amount of leaf chlorophyll. In the tree canopy, increasing I_D enhanced ζ_A through changes in M_A and N_A, whereas the direct effect of light was negative in both canopies. The overall correlation network between foliage structural and chemical traits and the relationships with I_D were significantly stronger in the tree canopy, suggesting limited re-acclimation potential in the mixed herbaceous canopy. Within-species acclimation patterns reflected the patterns within the main functional types. These data demonstrate that the relationships of current light availability vs. leaf dry mass per area, leaf nitrogen and chlorophyll contents, and chlorophyll to nitrogen ratio differ among multi-species herbaceous canopies and deciduous tree canopies due to contrasting canopy development.     

Effects of irradiance on growth,photosynthetic characteristics,and artemisinin content of <Emphasis Type="Italic">Artemisia annua</Emphasis> L.

With an increase in growth irradiance (from 15 to 100 % of full sunlight, I₁₅ to I₁₀₀), the maximum net photosynthetic rate (P _max), compensation (CI) and saturation irradiances of A. annua increased. At full sunlight, A. annua had a high capacity of photosynthesis, while at low irradiance it maintained a relatively high P _max with a low CI. The height and diameter growth, total and leaf biomass, and artemisinin content of A. annua decreased with the decrease in irradiance, which might be connected with lower photosynthesis at lower than at higher irradiance. Irradiances changed biomass allocations of A. annua. The leaf/total mass ratio of A. annua increased with decreasing irradiance, but the root/total mass ratio and root/above-ground mass generally increased with increasing irradiance. Thus A. annua can grow in both weak and full sunlight. However, high yield of biomass and artemisinin require cultivation in an open habitat with adequate sunshine.     

Three years of free-air CO2 enrichment (POPFACE) only slightly affect profiles of light and leaf characteristics in closed canopies of Populus

Modelling is used to predict long‐term forest responses to increased atmospheric CO₂ concentrations. Although productivity models are based on light intercepted by the canopy, very little experimental data are available for closed forest stands. Nevertheless, the relationships between light inside a canopy, leaf area, canopy structure, and individual leaf characteristics may be affected by elevated CO₂, affecting in turn carbon gain. Using a free‐air CO₂ enrichment (FACE) design in a high‐density plantation of Populus spp., we studied the effects of increased CO₂ concentrations on transmittance (τ) of photosynthetic photon flux density (Q_p), on ratios of red/far‐red light (R/FR), on leaf area index (LAI), on leaf inclination, on leaf chlorophyll (chl) and nitrogen (N) concentrations, and on specific leaf area (SLA) in the 2nd and 3rd years of treatment. Continuous measurements of τ were made in addition to canopy height profiles of light and leaf characteristics. Two years of Q_p measurements showed an average decrease of canopy transmittance in the FACE treatment, with very small differences at canopy closure. Results were explained by an unaffected LAI in closed canopies, without a FACE‐induced stimulation of relative crown depth. In agreement, leaf inclination and extinction coefficients for light were similar in control and FACE conditions. Ratios of R/FR were not significantly affected by the FACE treatment, neither were leaf characteristics, with the exception of leaf N, which allows speculation about N limitation. In general, treatment differences in canopy profiles resulted from an initial stimulation of height growth in the FACE treatment. P. × euramericana differed from P. alba and P. nigra, but species did not differ significantly in their response to the FACE treatment. By the time fast‐growing high‐density forest plantations have passed the exponential growth phase and reached canopy closure, the likely effects of elevated atmospheric CO₂ concentration on canopy architecture and absorption of Q_p are minor.     

Spatial distribution of leaf water-use efficiency and carbon isotope discrimination within an isolated tree crown

The spatial variations in the stable carbon isotope composition (δ¹³C) of air and leaves (total matter and soluble sugars) were quantified within the crown of a well‐watered, 20‐year‐old walnut tree growing in a low‐density orchard. The observed leaf carbon isotope discrimination (Δ) was compared with that computed by a three‐dimensional model simulating the intracanopy distribution of irradiance, transpiration and photosynthesis (previously parameterized and tested for the same tree canopy) coupled to a biophysically based model of carbon isotope discrimination. The importance of discrimination associated with CO₂ gradients encountered from the substomatal sites to the carboxylation sites was evaluated. We also assessed by simulation the effect of current irradiance on leaf gas exchange and the effect of long‐term acclimation of photosynthetic capacity and stomatal and internal conductances to light regime on intracanopy gradients in Δ. The main conclusions of this study are: (i) leaf Δ can exhibit important variations (5 and 8‰ in total leaf material and soluble sugars, respectively) along light gradients within the foliage of an isolated tree; (ii) internal conductance must be taken into account to adequately predict leaf Δ, and (iii) the spatial variations in Δ and water‐use efficiency resulted from the short‐term response of leaf gas exchange to variations in local irradiance and, to a much lesser extent, from the long‐term acclimation of leaf characteristics to the local light regime.     

Leaf and canopy responses of Lolium perenne to long-term elevated atmospheric carbon-dioxide concentration

The relationship between leaf photosynthetic capacity (p _{n, max}), net canopy CO₂- and H₂O-exchange rate (NCER and E _t, respectively) and canopy dry-matter production was examined in Lollium perenne L. cv. Vigor in ambient (363±30 l· l^-1) and elevated (631±43 l·l^-1) CO₂ concentrations. An open system for continuous and simultaneous regulation of atmospheric CO₂ concentration and NCER and E _t measurement was designed and used over an entire growth cycle to calculate a carbon and a water balance. While NCER_max of full-grown canopies was 49% higher at elevated CO₂ level, stimulation of p _n, max was only 46% (in spite of a 50% rise in one-sided stomatal resistance for water-vapour diffusion), clearly indicating the effect of a higher leaf-area index under high CO₂ (approx. 10% in one growing period examined). A larger amount of CO₂-deficient leaves resulted in higher canopy dark-respiration rates and higher canopy light compensation points. The structural component of the high-CO₂ effect was therefore a disadvantage at low irradiance, but a far greater benefit at high irradiance. Higher canopy darkrespiration rates under elevated CO₂ level and low irradiance during the growing period are the primary causes for the increase in dry-matter production (19%) being much lower than expected merely based on the NCER_max difference. While total water use was the same under high and low CO₂ levels, water-use efficiency increased 25% on the canopy level and 87% on a leaf basis. In the course of canopy development, allocation towards the root system became greater, while stimulation of shoot dry-matter accumulation was inversely affected. Over an entire growing season the root/shoot production ratio was 22% higher under high CO₂ concentration.Abbreviations and symbols C350 ambient CO₂, 363±30 l·l^-1 - C600 high CO₂, 631±43 l·l^-1 - c _a atmospheric CO₂ level - c _i CO₂ concentration in the intracellular spaces of the leaf - E_t canopy evapotranspiration - I _o canopy light compensation point - NCER canopy CO₂-exchange rate - p _n leaf photosynthetic rate - PPFD photosynthetic photon flux density - r _a leaf boundary-layer resistance - RD canopy dark-respiration rate - r _s stomatal resistance - WUE water use efficiency     

基于改进SW模型的千烟洲人工林蒸散组分拆分及其特征

蒸散组分拆分是准确评估陆地生态系统生产力以及估算水分利用效率的重要基础。利用改进后的Shuttleworth-Wallace模型,将蒸散拆分为植被蒸腾、土壤蒸发和冠层截留蒸发,并采用Monte Carlo随机参数化方案对模型参数进行优化。将模型与千烟洲亚热带人工针叶林站点的2011年涡度相关及小气候观测资料结合,对千烟洲人工林蒸散及其组分进行模拟。研究结果表明:半小时尺度上蒸散量模拟值与实测值的一致性在晴天和雨天都较高。半小时尺度上全年蒸散模拟值与实测值的决定系数、均方根误差和平均偏差为0.73、1.55 mmol m~(-2)s~(-1)和0.21 mmol m~(-2)s~(-1)。蒸散是该生态系统水分输出的最主要贡献项,占全年降水的80%。在蒸散中,植被蒸腾约占总蒸散量的85%,可推测2011年千烟洲人工林生态系统有较高的水分利用效率。该生态系统的蒸腾量季节变化明显,主要受饱和水汽压差和气温两种环境因素以及植被的叶面积指数影响且与三者均呈正相关;土壤蒸发约占总蒸散量的5%,季节变化平缓;模拟的冠层截留蒸发量约占总蒸散量的10%,季节变化大,与降水量呈正相关,与暴雨频次呈负相关,说明冠层无法有效截留强降水。该模型参数较少、时间分辨率高且可以有效模拟蒸散及其组分特征,是陆地生态系统水分循环过程研究有力的模型工具。     

Light Distribution in Discontinuous Canopies: Calculation of Leaf Areas and Canopy Volumes Above Defined 'Irradiance Contours' for use in Productivity Modelling

Plant canopies can be considered as assemblages of leaves, stemsand fruits growing in zones of differing irradiance demarcatedby contours of mean irradiance as measured on a horizontal surface. The following general equations have been derived to calculatethe leaf area (L_I) and the canopy volume (CV_I) in zones externalto any chosen contour of mean irradiance: (1) L_I = ((1nl)/(–K)(I–T_f) or leaf area index (LAI) if this is less (2) CV_I = L_I/(leaf area density m² m^–2), where I is the specified value of irradiance (horizontal surface)expressed as a decimal fraction of that above the canopy, Kis the appropriate extinction coefficient and T_f is the proportionof the total of available radiation which, if the canopy isdiscontinuous, would reach the ground by passing through gapsbetween the discrete canopy units. Where the canopy is continuousT_f is zero so expression (1) simplifies to L₁ = 1n I/–K(or LAI if this is less). For a range of model hedgerow orchards of varying dimensions,spacings and LAIs, it has been shown that the use of these equationsgives very similar results to those obtained by detailed calculationof light penetration. They therefore seem to be of potentialuse in calculating both potential dry-matter production by discontinuouscanopies of any type and, in the case of orchard fruit crops,the potential effect of changes in tree size, leaf area density,spacing etc. on the canopy volume in which irradiation is adequatefor fruit bud initiation and fruit colour development. light distribution, discontinuous canopy, irradiance contours, leaf area index, orchards