Plant competition for light analyzed with a multispecies canopy model

Summary Competition for light among species in a mixed canopy can be assessed quantitatively by a simulation model which evaluates the importance of different morphological and photosynthetic characteristics of each species. A model was developed that simulates how the foliage of all species attenuate radiation in the canopy and how much radiation is received by foliage of each species. The model can account for different kinds of foliage (leaf blades, stems, etc.) for each species. The photosynthesis and transpiration for sunlit and shaded foliage of each species is also computed for different layers in the canopy. The model is an extension of previously described single-species canopy photosynthesis simulation models. Model predictions of the fraction of foliage sunlit and interception of light by sunlit and shaded foliage for monoculture and mixed canopies of wheat (Triticum aestivum) and wild oat (Avena fatua) in the field compared very well with measured values. The model was used to calculate light interception and canopy photosynthesis for both species of wheat/wild oat mixtures grown under normal solar and enhanced ultraviolet-B (290–320 nm) radiation (UV-B) in a glasshouse experiment with no root competition. In these experiments, measurements showed that the mixtures receiving enhanced UV-B radiation had a greater proportion of the total foliage area composed of wheat compared to mixtures in the control treatments. The difference in species foliage area and its position in the canopy resulted in a calculated increase in the portion of total canopy radiation interception and photosynthesis by wheat. This, in turn, is consistent with greater canopy biomass of wheat reported in canopies irradiated with supplemental UV-B.     

Utilizing in situ directional hyperspectral measurements to validate bio-indicator simulations for a corn crop canopy

Two radiative transfer canopy models, SAIL and the two-layer Markov-Chain Canopy Reflectance Model (MCRM), were coupled with in situ leaf optical properties to simulate canopy-level spectral band ratio vegetation indices with the focus on the photochemical reflectance index in a cornfield. In situ hyperspectral measurements were made at both leaf and canopy levels. Leaf optical properties were obtained from both sunlit and shaded leaves. Canopy reflectance was acquired for eight different relative azimuth angles (ψ) at three different view zenith angles (θ_v), and later used to validate model outputs. Field observations of PRI for sunlit leaves exhibited lower values than shaded leaves, indicating higher light stress. Canopy PRI expressed obvious sensitivity to viewing geometry, as a function of both θ_v and ψ. Overall, simulations from MCRM exhibited better agreements with in situ values than SAIL. When using only sunlit leaves as input, the MCRM-simulated PRI values showed satisfactory correlation and RMSE, as compared to in situ values. However, the performance of the MCRM model was significantly improved after defining a lower canopy layer comprised of shaded leaves beneath the upper sunlit leaf layer. Four other widely used band ratio vegetation indices were also studied and compared with the PRI results. MCRM simulations were able to generate satisfactory simulations for these other four indices when using only sunlit leaves as input; but unlike PRI, adding shaded leaves did not improve the performance of MCRM. These results support the hypothesis that the PRI is sensitive to physiological dynamics while the others detect static factors related to canopy structure. Sensitivity analysis was performed on MCRM in order to better understand the effects of structure related parameters on the PRI simulations. LAI showed the most significant impact on MCRM-simulated PRI among the parameters studied. This research shows the importance of hyperspectral and narrow band sensor studies, and especially the necessity of including the green wavelengths (e.g., 531 nm) on satellites proposing to monitor carbon dynamics of terrestrial ecosystems.     

Microclimate and fluxes of water vapour, sensible heat and carbon dioxide in structurally differing subalpine plant communities in the Central Caucasus

Effects of canopy structure on microclimate, energy budget and CO₂ exchange were analysed in a pasture, two hay meadows, a tall herb community and a dwarf shrub community in the subalpine belt of the Central Caucasus. The results show that canopy structure exerts a marked influence on the distribution of photon flux density, temperature and canopy photosynthesis A_c. Three canopy types were distinguished. Type 1 (pasture) has a small LAI (leaf area index) and more than two-thirds of the phytomass is concentrated in the lowest few cm of the canopy, mainly as planophile leaves. This results in (1) a low degree of utilization of photosynthetic photon flux density (PPFD) by assimilatory plant components, (2) high leaf temperatures and a high soil heat flux during the phase of incoming radiation, and (3) a relatively low A_c/ LAI ratio. Type 2 (meadows), in spite of its erect leaves, which at high solar elevations permit light to penetrate to the lower canopy layers, is characterized by (1) marked effects of mutual shading in the lower canopy layers for most of the day, and thus (2) only slight variations in air and leaf temperatures and (3) a comparatively low A_c/LAI ratio. In canopies of type 3 (tall herb and dwarf shrub communities), there is a concentration of flat leaves in the upper layers. This results in (1) very good utilization of PPFD; (2) no strong fluctuations in canopy temperature as the flat leaves act as a buffer, reducing the amounts of incoming and outgoing radiation in lower canopy layers, and (3) high values of the A_c/LAI ratio. The energy budgets of the canopies investigated are governed not so much by their spatial structure, but rather indirectly by LAI and the degree of coupling of the canopy with the atmosphere.     

Spatial and temporal variability of canopy structure in a tropical moist forest

Tree fall gaps are widely considered to play a prominent role in the maintenance of species diversity, while the spatiotemporal variability of canopy structure within closed forest stands is largely ignored. In this study we examined the vertical and horizontal components of canopy structure and its seasonal variability in a tropical wet semideciduous rainforest in Panama. Leaf area indices (LAI) were derived from measurements of diffuse radiation and empirically-based leaf angle distribution by mathematical inversion of a light interception model. Vertical distribution of LAI was non-homogeneous with 50% of the leaf area being concentrated in the uppermost 5 m of the canopy. In the wet season, when foliage is most abundant, the horizontal distribution of LAI in a 2100 m² plot ranged widely from 3 to 8, with a mean of 5.41. Changes in mean LAI between wet and dry seasons were small but highly significant. While ca 40% of the area was not affected by local changes in LAI, sizeable small scale changes in LAI did occur between wet and dry season in some locations. Local changes in LAI ranged from –2.3 to 2.4. These changes resulted in a 50% or more increase in light reaching the forest floor at 29% of the measuring locations, and a doubling or more at 13% of the location. Our results imply that structural heterogeneity by simple tree fall gaps do not adequately describe the dynamics of forest canopies.     

Leaf nitrogen, photosynthesis, conductance and transpiration: scaling from leaves to canopies

A model is presented which solves simultaneously for leaf-scale stomatal conductance, CO₂ assimilation and the energy balance as a function of leaf position within canopies of well-watered vegetation. Fluxes and conductances were calculated separately for sunlit and shaded leaves. A linear dependence of photosynthetic capacity on leaf nitrogen content was assumed, while leaf nitrogen content and light intensity were assumed to decrease exponentially within canopies. Separate extinction coefficients were used for diffuse and direct beam radiation. An efficient Gaussian integration technique was used to compute fluxes and mean conductances for the canopy. The multilayer model synthesizes current knowledge of radiation penetration, leaf physiology and the physics of evaporation and provides insights into the response of whole canopies to multiple, interacting factors. The model was also used to explore sources of variation in the slopes of two simple parametric models (nitrogen- and light-use efficiency), and to set bounds on the magnitudes of the parameters. For canopies low in total N, daily assimilation rates are ～10% lower when leaf N is distributed uniformly than when the same total N is distributed according to the exponentially decreasing profile of absorbed radiation. However, gains are negligible for plants with high N concentrations. Canopy conductance, G_c should be calculated as G_c=Aσ(f_slg_sl+f_shg_sh), where Δ is leaf area index, f_si and f_sh are the fractions of sunlit and shaded leaves at each level, and g_si and g_sh are the corresponding stomatal conductances. Simple addition of conductances without this weighting causes errors in transpiration calculated using the ‘big-leaf’ version of the Penman-Monteith equation. Partitioning of available energy between sensible and latent heat is very responsive to the parameter describing the sensitivity of stomata to the atmospheric humidity deficit. This parameter also affects canopy conductance, but has a relatively small impact on canopy assimilation. Simple parametric models are useful for extrapolating understanding from small to large scales, but the complexity of real ecosystems is thus subsumed in unexplained variations in parameter values. Simulations with the multilayer model show that both nitrogen- and radiation-use efficiencies depend on plant nutritional status and the diffuse component of incident radiation, causing a 2- to 3-fold variation in these efficiencies.     

The phenology of leaf quality and its within‐canopy variation is essential for accurate modeling of photosynthesis in tropical evergreen forests

Leaf quantity (i.e., canopy leaf area index, LAI), quality (i.e., per‐area photosynthetic capacity), and longevity all influence the photosynthetic seasonality of tropical evergreen forests. However, these components of tropical leaf phenology are poorly represented in most terrestrial biosphere models (TBMs). Here, we explored alternative options for the representation of leaf phenology effects in TBMs that employ the Farquahar, von Caemmerer & Berry (FvCB) representation of CO₂ assimilation. We developed a two‐fraction leaf (sun and shade), two‐layer canopy (upper and lower) photosynthesis model to evaluate different modeling approaches and assessed three components of phenological variations (i.e., leaf quantity, quality, and within‐canopy variation in leaf longevity). Our model was driven by the prescribed seasonality of leaf quantity and quality derived from ground‐based measurements within an Amazonian evergreen forest. Modeled photosynthetic seasonality was not sensitive to leaf quantity, but was highly sensitive to leaf quality and its vertical distribution within the canopy, with markedly more sensitivity to upper canopy leaf quality. This is because light absorption in tropical canopies is near maximal for the entire year, implying that seasonal changes in LAI have little impact on total canopy light absorption; and because leaf quality has a greater effect on photosynthesis of sunlit leaves than light limited, shade leaves and sunlit foliage are more abundant in the upper canopy. Our two‐fraction leaf, two‐layer canopy model, which accounted for all three phenological components, was able to simulate photosynthetic seasonality, explaining ~90% of the average seasonal variation in eddy covariance‐derived CO₂ assimilation. This work identifies a parsimonious approach for representing tropical evergreen forest photosynthetic seasonality in TBMs that utilize the FvCB model of CO₂ assimilation and highlights the importance of incorporating more realistic phenological mechanisms in models that seek to improve the projection of future carbon dynamics in tropical evergreen forests.     

Angle distribution of foliage in individual Chamaecyparis obtusa canopies and effect of angle on diffuse light penetration

 The vertical distribution of foliage angle and area of three Chamaecyparis obtusa trees was determined by the triangle method, which calculates foliage geometry using measured coordinates of the leaf ”corners”, in a 43-year-old plantation in central Japan. Vertical distribution patterns of leaf area were different depending on tree size, but the boundary heights, which divide the canopy into sunlit and shaded parts, were similar in the three sample trees. The value of the average foliage angle [I(Z)] at a given depth (Z) from the tip of the stem decreased continually from the upper to lower layers within the canopy. The vertical patterns of changes in I(Z) were different among the three trees, but could be expressed by the following allometric equation as a function of depth.

Vegetation clumping modulates global photosynthesis through adjusting canopy light environment

The spatial dispersion of photoelements within a vegetation canopy, quantified by the clumping index (CI), directly regulates the within-canopy light environment and photosynthesis rate, but is not commonly implemented in terrestrial biosphere models to estimate the ecosystem carbon cycle. A few global CI products have been developed recently with remote sensing measurements, making it possible to examine the global impacts of CI. This study deployed CI in the radiative transfer scheme of the Community Land Model version 5 (CLM5) and used the revised CLM5 to quantitatively evaluate the extent to which CI can affect canopy absorbed radiation and gross primary production (GPP), and for the first time, considering the uncertainty and seasonal variation of CI with multiple remote sensing products. Compared to the results without considering the CI impact, the revised CLM5 estimated that sunlit canopy absorbed up to 9%–15% and 23%–34% less direct and diffuse radiation, respectively, while shaded canopy absorbed 3%–18% more diffuse radiation across different biome types. The CI impacts on canopy light conditions included changes in canopy light absorption, and sunlit–shaded leaf area fraction related to nitrogen distribution and thus the maximum rate of Rubisco carboxylase activity (V_cmax), which together decreased photosynthesis in sunlit canopy by 5.9–7.2 PgC year⁻¹ while enhanced photosynthesis by 6.9–8.2 PgC year⁻¹ in shaded canopy. With higher light use efficiency of shaded leaves, shaded canopy increased photosynthesis compensated and exceeded the lost photosynthesis in sunlit canopy, resulting in 1.0 ± 0.12 PgC year⁻¹ net increase in GPP. The uncertainty of GPP due to the different input CI datasets was much larger than that caused by CI seasonal variations, and was up to 50% of the magnitude of GPP interannual variations in the tropical regions. This study highlights the necessity of considering the impacts of CI and its uncertainty in terrestrial biosphere models.     

The effects of the spatial pattern of defoliation on regrowth of a tussock grass

Summary The spatial pattern of foliage removal from a tussock grass can influence regrowth through effects on daily carbon gain (CER_d). This field study examined the extent to which tussock photosynthetic responses to different defoliation patterns were associated with changes in whole-canopy attributes (e.g., foliage age structure, canopy light microclimate). During the spring growing season, 60% of the green foliage area was removed from individual Agropyron desertorum tussocks with scissors in different spatial patterns. These patterns represented extremes of defoliation patterns that might be inflicted by natural herbivores. Tussock photosynthesis (per unit foliage area) at high light (2000 mol photons m^–2 s^–1 between 400 and 700 nm; P₂₀₀₀) increased following clipping with all defoliation patterns. The increases in P₂₀₀₀ were greater when leaves were removed from low in the tussock (older leaves) than if leaves high in the canopy (younger leaves) were removed. These relative changes of P₂₀₀₀ among clipping patterns paralleled the responses of CER_d and regrowth from an earlier study. Furthermore, the changes in P₂₀₀₀ corresponded with increases in the proportion of foliage within the tussocks that was directly illuminated at midday. The greater photosynthesis of tussocks after lower-leaf removal was directly related to a higher proportion of younger foliage and a smaller fraction of foliage shaded within the tussock. In a dense canopy, such as these grass tussocks, the influence of defoliation on whole-canopy attributes may be of primary importance to whole-plant photosynthetic responses.     

On the nature of canopy illumination due to differences in elemental orientation and aggregation for radiative transfer

The nature of canopy radiative transfer mechanism (CRTM) describes the amount of beam penetration through a canopy and governs the nature of canopy illumination, i.e. the abundance of sunlit and shaded portions. Realistic representation of canopy illumination is critical for simulating various canopy biophysical processes associated with vegetated land surfaces. The adequate representation of CRTM can be attributed to the parameterizations of the two main canopy characteristics: the foliage projection (G-function) and the clumping effect (Ω function). Herein, using various types of G and Ω functions developed in a previous study, I tested 15 CRTM scenarios that combine different types of G and Ω functions to predict the dynamics of sunlit fraction (ε) of canopies having a wide range of plant area index (P_total) at various solar zenith angles (SZAs). It was observed that, for a given P_total, ε decreases as the SZA increases. However, ε significantly changed in accordance with the type of G and Ω functions used. Scenarios that employed random distribution of elements in space (S-4, S-9, and S-14) consistently returned larger ε values even at lower SZAs. This means that ignoring the clumping behavior of canopies could result in greater proportion of sunlit elements thereby reducing the beam penetration deeper into the canopy as opposed to those canopies where the elements are more aggregated. Beyond 70° SZA, almost all the scenarios returned similar ε values for a given P_total, which implied that the methods used is less sensitive at higher SZAs. The values of ε calculated by all the scenarios were significantly different from the S-6 (the ideal case). This observation highlights the importance of explicitly describing the G and Ω functions to adequately depict canopy illumination conditions.     

Regional climate modulates the canopy mosaic of favourable and risky microclimates for insects

1. One major gap in our ability to predict the impacts of climate change is a quantitative analysis of temperatures experienced by organisms under natural conditions. We developed a framework to describe and quantify the impacts of local climate on the mosaic of microclimates and physiological states of insects within tree canopies. This approach was applied to a leaf mining moth feeding on apple leaf tissues. 2. Canopy geometry was explicitly considered by mapping the 3D position and orientation of more than 26 000 leaves in an apple tree. Four published models for canopy radiation interception, energy budget of leaves and mines, body temperature and developmental rate of the leaf miner were integrated. Model predictions were compared with actual microclimate temperatures. The biophysical model accurately predicted temperature within mines at different positions within the tree crown. 3. Field temperature measurements indicated that leaf and mine temperature patterns differ according to the regional climatic conditions (cloudy or sunny) and depending on their location within the canopy. Mines in the sun can be warmer than those in the shade by several degrees and the heterogeneity of mine temperature was incremented by 120%, compared with that of leaf temperature. 4. The integrated model was used to explore the impact of both warm and exceptionally hot climatic conditions recorded during a heat wave on the microclimate heterogeneity at canopy scale. During warm conditions, larvae in sunlight-exposed mines experienced nearly optimal growth conditions compared with those within shaded mines. The developmental rate was increased by almost 50% in the sunny microhabitat compared with the shaded location. Larvae, however, experienced optimal temperatures for their development inside shaded mines during extreme climatic conditions, whereas larvae in exposed mines were overheating, leading to major risks of mortality. 5. Tree canopies act as both magnifiers and reducers of the climatic regime experienced in open air outside canopies. Favourable and risky spots within the canopy do change as a function of the climatic conditions at the regional scale. The shifting nature of the mosaic of suitable and risky habitats may explain the observed uniform distribution of leaf miners within tree canopies.     

RATP: a model for simulating the spatial distribution of radiation absorption, transpiration and photosynthesis within canopies: application to an isolated tree crown

The model RATP (radiation absorption, transpiration and photosynthesis) is presented. The model was designed to simulate the spatial distribution of radiation and leaf-gas exchanges within vegetation canopies as a function of canopy structure, canopy microclimate within the canopy and physical and physiological leaf properties. The model uses a three-dimensional (3D) representation of the canopy (i.e. an array of 3D cells, each characterized by a leaf area density). Radiation transfer is computed by a turbid medium analogy, transpiration by the leaf energy budget approach, and photosynthesis by the Farquhar model, each applied for sunlit and shaded leaves at the individual 3D cell-scale. The model typically operates at a 20–30 min time step. The RATP model was applied to an isolated, 20-year-old walnut tree grown in the field. The spatial distribution of wind speed, stomatal response to environmental variables, and light acclimation of leaf photosynthetic properties were taken into account. Model outputs were compared with data acquired in the field. The model was shown to simulate satisfactorily the intracrown distribution of radiation regime, transpiration and photosynthetic rates, at shoot or branch scales.     

Photosynthetic acclimation to elevated CO2 in a sunflower canopy

Sunflower canopies were grown in mesocosom gas exchange chambers at ambient and elevated CO2 concentrations (360 and 700 ppm) and leaf photosynthetic capacities measured at several depths within each canopy. Elevated [CO2] had little effect on whole-canopy photosynthetic capacity and total leaf area, but had marked effects on the distribution of photosynthetic capacity and leaf area within the canopy. Elevated [CO2] did not significantly reduce the photosynthetic capacities per unit leaf area of young leaves at the top of the canopy, but it did reduce the photosynthetic capacities of older leaves by as much as 40%. This effect was not dependent on the canopy light environment since elevated [CO2] also reduced the photosynthetic capacities of older leaves exposed to full sun on the south edge of the canopy. In addition to the effects on leaf photosynthetic capacity, elevated [CO2] shifted the distribution of leaf area within the canopy so that more leaf area was concentrated near the top of the canopy. This change resulted in as much as a 50% reduction in photon flux density in the upper portions of the elevated [CO2] canopy relative to the ambient [CO2] canopy, even though there was no significant difference in the total canopy leaf area. This reduction in PFD appeared to account for leaf carbohydrate contents that were actually lower for many of the shaded leaves in the elevated as opposed to the ambient [CO2] canopy. Photosynthetic capacities were not significantly correlated with any of the individual leaf carbohydrate contents. However, there was a strong negative correlation between photosynthetic capacity and the ratio of hexose sugars to sucrose, consistent with the hypothesis that sucrose cycling is a component of the biochemical signalling pathway controlling photosynthetic acclimation to elevated [CO2].     

Leaf traits and associated ecosystem characteristics across subtropical and timberline forests in the Gongga Mountains,Eastern Tibetan Plateau

Knowledge of how leaf characteristics might be used to deduce information on ecosystem functioning and how this scaling task could be done is limited. In this study, we present field data for leaf lifespan, specific leaf area (SLA) and mass and area-based leaf nitrogen concentrations (N_mass, N_area) of dominant tree species and the associated stand foliage N-pool, leaf area index (LAI), root biomass, aboveground biomass, net primary productivity (NPP) and soil available-N content in six undisturbed forest plots along subtropical to timberline gradients on the eastern slope of the Gongga Mountains. We developed a methodology to calculate the whole-canopy mean leaf traits to include all tree species (groups) in each of the six plots through a series of weighted averages scaled up from leaf-level measurements. These defined whole-canopy mean leaf traits were equivalent to the traits of a leaf in regard to their interrelationships and altitudinal trends, but were more useful for large-scale pattern analysis of ecosystem structure and function. The whole-canopy mean leaf lifespan and leaf N_mass mainly showed significant relationships with stand foliage N-pool, NPP, LAI and root biomass. In general, as elevation increased, the whole-canopy mean leaf lifespan and leaf N_area and stand LAI and foliage N-pool increased to their maximum, whereas the whole-canopy mean SLA and leaf N_mass and stand NPP and root biomass decreased from their maximum. The whole-canopy mean leaf lifespan and stand foliage N-pool both converged towards threshold-like logistic relationships with annual mean temperature and soil available-N variables. Our results are further supported by additional literature data in the Americas and eastern China.     

Combining thermal and visible imagery for estimating canopy temperature and identifying plant stress

Thermal imaging is a potential tool for estimating plant temperature, which can be used as an indicator of stomatal closure and water deficit stress. In this study, a new method for processing and analysing thermal images was developed. By using remote sensing software, the information from thermal and visible images was combined, the images were classified to identify leaf area and sunlit and shaded parts of the canopy, and the temperature statistics for specific canopy components were calculated. The method was applied to data from a greenhouse water-stress experiment of Vicia faba L. and to field data for Vitis vinifera L. Vaseline-covered and water-sprayed plants were used as dry and wet references, respectively, and two thermal indices, based on temperature differences between the canopy and reference surfaces, were calculated for single Vicia faba plants. The thermal indices were compared with measured stomatal conductance. The temperature distributions of sunlit and shaded leaf area of Vitis vinifera canopies from natural rainfall and irrigation treatments were compared. The present method provides two major improvements compared with earlier methods for calculating thermal indices. First, it allows more accurate estimation of the indices, which are consequently more closely related to stomatal conductance. Second, it gives more accurate estimates of the temperature distribution of the shaded and sunlit parts of canopy, and, unlike the earlier methods, makes it possible to quantify the relationship between temperature variation and stomatal conductance.     

Scaling CO2-photosynthesis relationships from the leaf to the canopy

Responses of individual leaves to short-term changes in CO₂ partial pressure have been relatively well studied. Whole-plant and plant community responses to elevated CO₂ are less well understood and scaling up from leaves to canopies will be complicated if feedbacks at the small scale differ from feedbacks at the large scale. Mathematical models of leaf, canopy, and ecosystem processes are important tools in the study of effects on plants and ecosystems of global environmental change, and in particular increasing atmospheric CO₂, and might be used to scale from leaves to canopies. Models are also important in assessing effects of the biosphere on the atmosphere. Presently, multilayer and big leaf models of canopy photosynthesis and energy exchange exist. Big leaf models — which are advocated here as being applicable to the evaluation of impacts of global change on the biosphere — simplify much of the underlying leaf-level physics, physiology, and biochemistry, yet can retain the important features of plant-environment interactions with respect to leaf CO₂ exchange processes and are able to make useful, quantitative predictions of canopy and community responses to environmental change. The basis of some big leaf models of photosynthesis, including a new model described herein, is that photosynthetic capacity and activity are scaled vertically within a canopy (by plants themselves) to match approximately the vertical profile of PPFD. The new big leaf model combines physically based models of leaf and canopy level transport processes with a biochemically based model of CO₂ assimilation. Predictions made by the model are consistent with canopy CO₂ exchange measurements, although a need exists for further testing of this and other canopy physiology models with independent measurements of canopy mass and energy exchange at the time scale of 1 h or less.Abbreviations LAI leaf area index - NIR near infrared (700–3000 nm) radiation - PAR photosynthetically active (400–700 nm) radiation - PI photosynthetic irradiance (400–700 nm) - PPFD photosynthetic photon flux area density (400–700 nm) - PS I Photosystem I - PS II Photosystem II - Rubisco ribulose-1,5-bisphosphate carboxylase/oxygenase - RuP₂ ribulose-1,5-bisphosphate     

Effects of CO2 elevation on canopy development in the stands of two co-occurring annuals

Elevated CO₂ may increase dry mass production of canopies directly through increasing net assimilation rate of leaves and also indirectly through increasing leaf area index (LAI). We studied the effects of CO₂ elevation on canopy productivity and development in monospecific and mixed (1:1) stands of two co-occurring C₃ annual species, Abutilon theophrasti, and Ambrosia artemisiifolia. The stands were established in the glasshouse with two CO₂ levels (360 and 700 l/l) under natural light conditions. The planting density was 100 per m² and LAI increased up to 2.6 in 53 days of growth. Root competition was excluded by growing each plant in an individual pot. However, interference was apparent in the amount of photons absorbed by the plants and in photon absorption per unit leaf area. Greater photon absorption by Abutilon in the mixed stand was due to different canopy structures: Abutilon distributed leaves in the upper layers in the canopy while Ambrosia distributed leaves more to the lower layers. CO₂ elevation did not affect the relative performance and light interception of the two species in mixed stands. Total aboveground dry mass was significantly increased with CO₂ elevation, while no significant effects on leaf area development were observed. CO₂ elevation increased dry mass production by 30–50%, which was mediated by 35–38% increase in the net assimilation rate (NAR) and 37–60% increase in the nitrogen use efficiency (NUE, net assimilation rate per unit leaf nitrogen). Since there was a strong overall correlation between LAI and aboveground nitrogen and no significant difference was found in the regression of LAI against aboveground nitrogen between the two CO₂ levels, we hypothesized that leaf area development was controlled by the amount of nitrogen taken up from the soil. This hypothesis suggests that the increased LAI with CO₂ elevation observed by several authors might be due to increased uptake of nitrogen with increased root growth.     

Scaling Up Stomatal Conductance from Leaf to Canopy Using a Dual-Leaf Model for Estimating Crop Evapotranspiration

The dual-source Shuttleworth-Wallace model has been widely used to estimate and partition crop evapotranspiration (λET). Canopy stomatal conductance (G_sc), an essential parameter of the model, is often calculated by scaling up leaf stomatal conductance, considering the canopy as one single leaf in a so-called “big-leaf” model. However, G_sc can be overestimated or underestimated depending on leaf area index level in the big-leaf model, due to a non-linear stomatal response to light. A dual-leaf model, scaling up G_sc from leaf to canopy, was developed in this study. The non-linear stomata-light relationship was incorporated by dividing the canopy into sunlit and shaded fractions and calculating each fraction separately according to absorbed irradiances. The model includes: (1) the absorbed irradiance, determined by separately integrating the sunlit and shaded leaves with consideration of both beam and diffuse radiation; (2) leaf area for the sunlit and shaded fractions; and (3) a leaf conductance model that accounts for the response of stomata to PAR, vapor pressure deficit and available soil water. In contrast to the significant errors of G_sc in the big-leaf model, the predicted G_sc using the dual-leaf model had a high degree of data-model agreement; the slope of the linear regression between daytime predictions and measurements was 1.01 (R² = 0.98), with RMSE of 0.6120 mm s⁻¹ for four clear-sky days in different growth stages. The estimates of half-hourly λET using the dual-source dual-leaf model (DSDL) agreed well with measurements and the error was within 5% during two growing seasons of maize with differing hydrometeorological and management strategies. Moreover, the estimates of soil evaporation using the DSDL model closely matched actual measurements. Our results indicate that the DSDL model can produce more accurate estimation of G_sc and λET, compared to the big-leaf model, and thus is an effective alternative approach for estimating and partitioning λET.     

Patterns of cytokinins and leaf gas exchange among canopy layers of a mature sugar maple (<Emphasis Type="Italic">Acer saccharum</Emphasis>) stand

Leaf cytokinins (CKs) were profiled within four locations throughout the inner and outer layers of a mature sugar maple (Acer saccharum) canopy. Leaf CK was associated with leaf gas exchange activity and some corresponding microclimate variables. Both inner and outer layers in the upper canopy had higher concentrations of leaf CKs than the lower canopy layers and the difference was comprised primarily by riboside forms of CK. Transpiration (E) showed a similar pattern to leaf CK content, with significantly higher rates in the upper canopy. There was, however, no clear pattern discernable in stomatal conductance (gs), other than it tended to be higher in the outer canopy layers. The upper/outer canopy showed a significantly different environment than all other canopy positions with higher photosynthetically active radiation (PAR), ultra-violet light (UV-B) and leaf temperature. Simple linear regression analysis showed that the nucleotide CK group (including iPNT, cis- and trans-[9RMP]Z, [9RMP]DZ) was positively related to PAR. Exogenous applications of benzylaminopurine (BAP), showed that low concentrations of BAP reduced E and g _s, and indicated that CK may help regulate stomatal aperture. The similar patterns in E and CK content suggest that CKs and leaf gas exchange are functionally connected.     

Effects of drought on mesophyll conductance and photosynthetic limitations at different tree canopy layers

In recent years, many studies have focused on the limiting role of mesophyll conductance (g_m) to photosynthesis (A_n) under water stress, but no studies have examined the effect of drought on g_m through the forest canopy. We investigated limitations to A_n on leaves at different heights in a mixed adult stand of sessile oak (Quercus petraea) and beech (Fagus sylvatica) trees during a moderately dry summer. Moderate drought decreased A_n of top and lowest beech canopy leaves much more than in leaves located in the mid canopy; whereas in oak, A_n of the lower canopy was decreased more than in sunlit leaves. The decrease of A_n was probably not due to leaf‐level biochemistry given that V_Cmax was generally unaffected by drought. The reduction in A_n was instead associated with reduction in stomatal and mesophyll conductances. Drought‐induced increases in stomatal limitations were largest in leaves from the top canopy, whereas drought‐induced increases in mesophyll limitations were largest in leaves from the lowest canopy. Sensitivity analysis highlighted the need to decompose the canopy into different leaf layers and to incorporate the limitation imposed by g_m when assessing the impact of drought on the gas exchange of tree canopies.