Modelling canopy CO2 fluxes: are ‘big‐leaf’ simplifications justified?

• 1 The ‘big‐leaf’ approach to calculating the carbon balance of plant canopies is evaluated for inclusion in the ETEMA model framework. This approach assumes that canopy carbon fluxes have the same relative responses to the environment as any single leaf, and that the scaling from leaf to canopy is therefore linear.
• 2 A series of model simulations was performed with two models of leaf photosynthesis, three distributions of canopy nitrogen, and two levels of canopy radiation detail. Leaf‐ and canopy‐level responses to light and nitrogen, both as instantaneous rates and daily integrals, are presented.
• 3 Observed leaf nitrogen contents of unshaded leaves are over 40% lower than the big‐leaf approach requires. Scaling from these leaves to the canopy using the big‐leaf approach may underestimate canopy photosynthesis by ~20%. A leaf photosynthesis model that treats within‐leaf light extinction displays characteristics that contradict the big‐leaf theory. Observed distributions of canopy nitrogen are closer to those required to optimize this model than the homogeneous model used in the big‐leaf approach.
• 4 It is theoretically consistent to use the big‐leaf approach with the homogeneous photosynthesis model to estimate canopy carbon fluxes if canopy nitrogen and leaf area are known and if the distribution of nitrogen is assumed optimal. However, real nitrogen profiles are not optimal for this photosynthesis model, and caution is necessary in using the big‐leaf approach to scale satellite estimates of leaf physiology to canopies. Accurate prediction of canopy carbon fluxes requires canopy nitrogen, leaf area, declining nitrogen with canopy depth, the heterogeneous model of leaf photosynthesis and the separation of sunlit and shaded leaves. The exact nitrogen profile is not critical, but realistic distributions can be predicted using a simple model of canopy nitrogen allocation.

     

Plant ecophysiological processes in spectral profiles: perspective from a deciduous broadleaf forest

The need for progress in satellite remote sensing of terrestrial ecosystems is intensifying under climate change. Further progress in Earth observations of photosynthetic activity and primary production from local to global scales is fundamental to the analysis of the current status and changes in the photosynthetic productivity of terrestrial ecosystems. In this paper, we review plant ecophysiological processes affecting optical properties of the forest canopy which can be measured with optical remote sensing by Earth-observation satellites. Spectral reflectance measured by optical remote sensing is utilized to estimate the temporal and spatial variations in the canopy structure and primary productivity. Optical information reflects the physical characteristics of the targeted vegetation; to use this information efficiently, mechanistic understanding of the basic consequences of plant ecophysiological and optical properties is essential over broad scales, from single leaf to canopy and landscape. In theory, canopy spectral reflectance is regulated by leaf optical properties (reflectance and transmittance spectra) and canopy structure (geometrical distributions of leaf area and angle). In a deciduous broadleaf forest, our measurements and modeling analysis of leaf-level characteristics showed that seasonal changes in chlorophyll content and mesophyll structure of deciduous tree species lead to a seasonal change in leaf optical properties. The canopy reflectance spectrum of the deciduous forest also changes with season. In particular, canopy reflectance in the green region showed a unique pattern in the early growing season: green reflectance increased rapidly after leaf emergence and decreased rapidly after canopy closure. Our model simulation showed that the seasonal change in the leaf optical properties and leaf area index caused this pattern. Based on this understanding we discuss how we can gain ecophysiological information from satellite images at the landscape level. Finally, we discuss the challenges and opportunities of ecophysiological remote sensing by satellites.

     

Vegetation–Climate Interactions among Native and Invasive Species in Hawaiian Rainforest

We compiled a time series of Earth Observing-1 Hyperion satellite observations with field measurements to compare the structural, biochemical, and physiological characteristics of an invasive nitrogen-fixing tree Myrica faya and native Metrosideros polymorpha in montane rainforests in Hawai’i. Satellite-based canopy water measurements closely tracked variations in leaf area index, and the remotely sensed photochemical and carotenoid reflectance indices (PRI, CRI) indicated variations in upper-canopy leaf chlorophyll and carotenoid content during a climatological transition. The PRI and CRI were related to differences in light-use efficiency of each species, as indicated by field measurements of leaf electron transport rate. The suite of hyperspectral metrics indicated maximum differences in the structure, biochemistry, and physiology of Myrica and Metrosideros when canopy vapor pressure deficit was high during hotter and drier periods. These satellite data, combined with the Carnegue-Ames-Stanford Approach (CASA) carbon cycle model, suggested that Myrica growth rates were 16–44% higher than Metrosideros, with relative differences between species closely linked to climate conditions. The satellite hyperspectral data identified the basic biological mechanisms favoring the spread of an introduced tree, and provided a more detailed understanding of how vegetation–climate interactions affect the time course of plant growth with respect to the invasion process.     

Canopy photosynthetic production in a Japanese larch forest. II. Estimation of the canopy photosynthetic production

Seasonal changes and yearly gross canopy photosynthetic production were estimated for an 18 year old Japanese larch (Larix leptolepis) forest between 1982 and 1984. A canopy photosynthesis model was applied for the estimation, which took into account the effect of light interception by the non-photosynthetic organs. Seasonal changes in photosynthetic ability, amount of canopy leaf area and light environment within the canopy were also taken into account. Amount of leaf area was estimated by the leaf area growth of a single leaf. The change of light environment within the canopy during the growing season was estimated with a light penetration model and the leaf increment within the canopy. Canopy respiration and surplus production were calculated as seasonal and yearly values for the three years studied. Mean yearly estimates of canopy photosynthesis, canopy respiration and surplus production were 37, 13 and 23 tCO₂ ha⁻¹ year⁻¹, respectively. Vertical trend, seasonal changes and yearly values of the estimates were analyzed in relation to environmental and stand factors.     

Modeling the date of leaf appearance in low-arctic tundra

One of the reported changes of arctic ecosystems in response to warming climate is the advance of the leaf appearance in spring. Such phenological changes play a role in the structural changes within tundra ecosystem communities. Recently, we developed a model that estimates the leaf appearance date for deciduous trees in taiga. We apply this model to the whole low-arctic tundra, and we compare the simulated green-up dates with the green-up dates obtained from satellite observations and to in situ measurements of deciduous shrub leaf appearance. The model, although calibrated for taiga, performs remarkably well in tundra, with root mean square error ranging between 4 and 8 days for most of the tundra region, the same order as in taiga regions. The results seem to indicate that air temperature is the main factor controlling spring leaf phenology in tundra, just as in taiga, although these results do not permit us to reject soil temperature as the main trigger for leaf appearance in tundra. Because our model performs in tundra as well as in taiga, it can be used across the ecotone, and during a northward migration of the species from the taiga to the low-arctic region. The leaf appearance model and the satellite observations reveal that leaf appearance has tended to occur earlier by approximately 10 days both in Alaska since 1975, and in west Siberian tundra since 1965.     

Linking drought legacy effects across scales: From leaves to tree rings to ecosystems

Severe drought can cause lagged effects on tree physiology that negatively impact forest functioning for years. These “drought legacy effects” have been widely documented in tree‐ring records and could have important implications for our understanding of broader scale forest carbon cycling. However, legacy effects in tree‐ring increments may be decoupled from ecosystem fluxes due to (a) postdrought alterations in carbon allocation patterns; (b) temporal asynchrony between radial growth and carbon uptake; and (c) dendrochronological sampling biases. In order to link legacy effects from tree rings to whole forests, we leveraged a rich dataset from a Midwestern US forest that was severely impacted by a drought in 2012. At this site, we compiled tree‐ring records, leaf‐level gas exchange, eddy flux measurements, dendrometer band data, and satellite remote sensing estimates of greenness and leaf area before, during, and after the 2012 drought. After accounting for the relative abundance of tree species in the stand, we estimate that legacy effects led to ~10% reductions in tree‐ring width increments in the year following the severe drought. Despite this stand‐scale reduction in radial growth, we found that leaf‐level photosynthesis, gross primary productivity (GPP), and vegetation greenness were not suppressed in the year following the 2012 drought. Neither temporal asynchrony between radial growth and carbon uptake nor sampling biases could explain our observations of legacy effects in tree rings but not in GPP. Instead, elevated leaf‐level photosynthesis co‐occurred with reduced leaf area in early 2013, indicating that resources may have been allocated away from radial growth in conjunction with postdrought upregulation of photosynthesis and repair of canopy damage. Collectively, our results indicate that tree‐ring legacy effects were not observed in other canopy processes, and that postdrought canopy allocation could be an important mechanism that decouples tree‐ring signals from GPP.     

Tracking plant physiological properties from multi-angular tower-based remote sensing

Imaging spectroscopy is a powerful technique for monitoring the biochemical constituents of vegetation and is critical for understanding the fluxes of carbon and water between the land surface and the atmosphere. However, spectral observations are subject to the sun–observer geometry and canopy structure which impose confounding effects on spectral estimates of leaf pigments. For instance, the sun–observer geometry influences the spectral brightness measured by the sensor. Likewise, when considering pigment distribution at the stand level scale, the pigment content observed from single view angles may not necessarily be representative of stand-level conditions as some constituents vary as a function of the degree of leaf illumination and are therefore not isotropic. As an alternative to mono-angle observations, multi-angular remote sensing can describe the anisotropy of surface reflectance and yield accurate information on canopy structure. These observations can also be used to describe the bi-directional reflectance distribution which then allows the modeling of reflectance independently of the observation geometry. In this paper, we demonstrate a method for estimating pigment contents of chlorophyll and carotenoids continuously over a year from tower-based, multi-angular spectro-radiometer observations. Estimates of chlorophyll and carotenoid content were derived at two flux-tower sites in western Canada. Pigment contents derived from inversion of a CR model (PROSAIL) compared well to those estimated using a semi-analytical approach (r ² = 0.90 and r ² = 0.69, P < 0.05 for both sites, respectively). Analysis of the seasonal dynamics indicated that net ecosystem productivity was strongly related to total canopy chlorophyll content at the deciduous site (r ² = 0.70, P < 0.001), but not at the coniferous site. Similarly, spectral estimates of photosynthetic light-use efficiency showed strong seasonal patterns in the deciduous stand, but not in conifers. We conclude that multi-angular, spectral observations can play a key role in explaining seasonal dynamics of fluxes of carbon and water and provide a valuable addition to flux-tower-based networks.     

棉花叶片氮含量的空间分布与光合特性

在棉花生长旺季,将冠层按高度分多层测定了田间叶片含氮量和叶片净光合速率对光合有效辐射通量密度的响应(光响应曲线,Pn-PPFD response curve)及相应的生物指标.结果表明,各层叶片氮含量与光合作用关系密切,各层平均值大小依次为上层＞中层＞下层,对应层叶片的最大净光合速率Pmax、表观暗呼吸速率Rd、光补偿点LCP及光饱和点LSP均从上到下依次递减,与氮含量分布一致,而表观光合量子效率AQY则略有不同;氮含量的指数衰减系数 kn =0.762(R2=0.593),根据观测结果,棉田叶片氮含量(N)空间分布可以用相对累积叶面积指数(Lc/Lt)为自变量的指数方程来模拟,从而为建立光合作用机理模型与进行生产力奠定基础.     

Canopy structure in savannas along a moisture gradient on Kalahari sands

Measurements of tree canopy architecture were made at six savanna sites on deep, sandy soils, along a gradient of increasing aridity. There was substantial variation in the leaf area estimated within each site, using the same sample frame, but different measurement techniques. The trends in canopy properties in relation to the aridity gradient were consistent, regardless of the technique used for estimating the properties. The effective plant area index for the tree canopy (the sum of the stem area index and the leaf area index (LAI)) declined from around 2 to around 0.8 m² m^?2 over a gradient of mean annual rainfall from 1000 to 350 mm. Stems contributed 2–5% of the tree canopy plant area index. Since the tree canopy cover decreased from 50% to 20% over this aridity range, the leaf area index within the area covered by tree canopies remained fairly constant at 3–4 m² m^?2. Tree leaves tended from a horizontal orientation to a more random orientation as the aridity increased. On the same gradient, the leaf minor axis dimension decreased from around 30 mm to around 3 mm, and the mean specific leaf area decreased from 14 to 5 m² kgha^?1. There was good agreement between LAI observed in the field using a line ceptometer and the LAI inferred by the MODIS sensor on the Terra satellite platform, 2 months later in the same season.     

Estimating canopy conductance to ozone uptake from observations of evapotranspiration at the canopy scale and at the leaf scale*

Stomatal uptake by vegetation is often the major sink for the destruction of tropospheric ozone. Using data obtained during the summer of 1991 at a grape vineyard and a cotton field in the San Joaquin Valley of California, we compare canopy (stomatal) conductances to ozone estimated (1) from eddy covariance ozone flux data (2) from eddy covariance evapotranspiration data and (3) by scaling leaf trans pi rational conductance to the canopy level using a canopy radiative transfer model. These simultaneous data, obtained at two levels of biological organization and for two trace gases, allow us to contrast the pathways for canopy-atmosphere exchange of water vapour and ozone, to evaluate limitations to scaling from leaf to canopy, and to predict ozone uptake parameters from those governing transpiration. At the vineyard site the eddy covariance ozone results underestimate the ET-based (eddy covariance and leaf scaling) approaches between 25% and 36%. At the cotton site the ozone-based results overestimate the ET-based approaches between 9% and 62%. A number of modelling and measurement uncertainties are of appropriate magnitude to reconcile these estimates. Some of the possible causes for these discrepancies that are discussed include NO effects, mesophyll resistances to ozone uptake and flaws in the K-theory (first-order closure) approach on which the canopy-scale analysis is based. Nevertheless, both canopy and single leaf measurements of conductance for water vapour provide acceptable estimates of conductance for ozone, but further experiments in which all are measured simultaneously are suggested.     

Estimates of tree canopy loss as a result of Cyclone Monica,in the Magela Creek catchment northern Australia

Abstract Severe tropical Cyclone Monica impacted the coast of northern Australia in April 2006 with estimated maximum wind gusts of 360 km h^?1. It rapidly moved inland losing intensity and passed over the town of Jabiru as a category 2 system, with maximum wind gusts recorded at 135 km h^?1. The cyclone had a significant impact on the landscapes within the Alligator Rivers Region and significant windthrow of trees occurred. This paper describes the level of impact that category 2 level winds had on tree canopy loss 10 days after cyclone and then again 1 year later. Recovery was assessed using multispectral satellite imagery in sub‐catchments of the Magela Creek catchments. A non‐linear relationship was fitted between a modified vegetation index (derived from Landsat TM5 satellite data) and percentage tree canopy cover (measured from very high resolution QuickBird satellite data). The results of the non‐linear relationship, used to estimate percentage canopy cover, indicate that 10 days after cyclone, there was significant disturbance to tree canopy. However, data 1 year after cyclone show that recovery of canopy across the studied catchments varied between 8% and 19% of the percentage canopy cover that remained after the initial impact of the cyclone. Further analysis in the three sub‐catchments using Geographical Information System showed that proportionally, riparian zones and inundated areas in each of the sub‐catchments suffered greater loss of tree canopy cover compared with upland areas.     

Measured and modelled leaf and stand‐scale productivity across a soil moisture gradient and a severe drought

Environmental controls on carbon dynamics operate at a range of interacting scales from the leaf to landscape. The key questions of this study addressed the influence of water and nitrogen (N) availability on Pinus palustris (Mill.) physiology and primary productivity across leaf and canopy scales, linking the soil‐plant‐atmosphere (SPA) model to leaf and stand‐scale flux and leaf trait/canopy data. We present previously unreported ecophysiological parameters (e.g. V_cmax and J_max) for P. palustris and the first modelled estimates of its annual gross primary productivity (GPP) across xeric and mesic sites and under extreme drought. Annual mesic site P. palustris GPP was ～23% greater than at the xeric site. However, at the leaf level, xeric trees had higher net photosynthetic rates, and water and light use efficiency. At the canopy scale, GPP was limited by light interception (canopy level), but co‐limited by nitrogen and water at the leaf level. Contrary to expectations, the impacts of an intense growing season drought were greater at the mesic site. Modelling indicated a 10% greater decrease in mesic GPP compared with the xeric site. Xeric P. palustris trees exhibited drought‐tolerant behaviour that contrasted with mesic trees' drought‐avoidance behaviour.     

Growth Response of a Chrysanthemum Crop to the Environment. III, Effects of Radiation and Temperature on Dry Matter Partitioning and Photosynthesis

Vegetative crops of chrysanthemum were grown for 5 or 6 weekperiods in daylit assimilation chambers. Crop responses to differentradiation levels and temperatures were analysed into effectson dry matter partitioning, specific leaf area, leaf photosynthesisand canopy light interception. The percentage of newly formed dry matter partitioned to theleaves was almost constant, although with increasing radiationor decreasing temperature, a greater percentage of dry matterwas partitioned to stem tissue at the expense of root tissue.There was a positive correlation between the percentage of drymatter in shoot material and the overall carbon: dry matterratio. Canopy photosynthesis was analysed assuming identical behaviourfor all leaves in the crop. Leaf photochemical efficiency wasonly slightly affected by crop environment. The rate of grossphotosynthesis per unit leaf area at light saturation, P_A (max),increased with increasing radiation integral, but the same parameterexpressed per unit leaf dry matter, P_w (max) was almost unaffectedby growth radiation. In contrast, P_A (max) was hardly affectedby temperature but P_w (max) increased with increasing growthtemperature. This was because specific leaf area decreased withdecreasing temperature and increased with decreasing radiation.There was a positive correlation between canopy respirationintegral and photosynthesis integral, and despite a four-foldchange in crop mass during the experiments, the maintenancecomponent of canopy respiration remained small and constant. Canopy extinction coefficient showed no consistent variationwith radiation integral but was negatively correlated with temperature.This decrease in the efficiency of the canopy at interceptingradiation exactly cancelled the increase in specific carbonassimilation rate that occurred with increasing growth temperature,giving a growth rate depending solely on the incident lightlevel. Chrysanthemum, dry matter partitioning, photosynthesis, specific leaf area     

Stomatal control of transpiration from a developing sugarcane canopy

Abstract. Stomatal conductance of single leaves and transpiration from an entire sugarcane (Saccharum spp. hybrid) canopy were measured simultaneously using independent techniques. Stomatal and environmental controls of transpiration were assessed at three stages of canopy development, corresponding to leaf area indices (L) of 2.2, 3.6 and 5.6. Leaf and canopy boundary layers impeded transport of transpired water vapour away from the canopy, causing humidity around the leaves to find its own value through local equilibration rather than a value determined by the humidity of the bulk air mass above the canopy. This tended to uncouple transpiration from direct stomatal control, so that transpiration predicted from measurement of stomatal conductance and leaf-to-air vapour pressure differences was increasingly overestimated as the reference point for ambient vapour pressure measurement was moved farther from the leaf and into the bulk air. The partitioning of control between net radiation and stomata was expressed as a dimensionless decoupling coefficent ranging from zero to 1.0. When the stomatal aperture was near its maximum this coefficient was approximately 0.9, indicating that small reductions in stomatal aperture would have had little effect on canopy transpiration. Maximum rates of transpiration were, however, limited by large adjustments in maximum stomatal conductance during canopy development. The product of maximum stomatal conductance and L. a potential total canopy conductance in the absence of boundary layer effects, remained constant as L increased. Similarly, maximum canopy conductance, derived from independent micrometeorological measurements, also remained constant over this period. Calculations indicated that combined leaf and canopy boundary layer conductance decreased with increasing L such that the ratio of boundary layer conductance to maximum stomatal conductance remained nearly constant at approximately 0.5. These observations indicated that stomata adjusted to maintain both transpiration and the degree of stomatal control of transpiration constant as canopy development proceeded.     

Variation in potential for isoprene emissions among Neotropical forest sites

As part of the Large Scale Biosphere–Atmosphere Experiment in Amazônia (LBA), we have developed a bottom‐up approach for estimating canopy‐scale fluxes of isoprene. Estimating isoprene fluxes for a given forest ecosystem requires knowledge of foliar biomass, segregated by species, and the isoprene emission characteristics of the individual tree species comprising the forest. In this study, approximately 38% of 125 tree species examined at six sites in the Brazilian Amazon emitted isoprene. Given logistical difficulties and extremely high species diversity, it was possible to screen only a small percentage of tree species, and we propose a protocol for estimating the emission capacity of unmeasured taxa using a taxonomic approach, in which we assign to an unmeasured genus a value based on the percentage of genera within its plant family which have been shown to emit isoprene. Combining this information with data obtained from 14 tree censuses at four Neotropical forest sites, we have estimated the percentage of isoprene‐emitting biomass at each site. The relative contribution of each genus of tree is estimated as the basal area of all trees of that genus divided by the total basal area of the plot. Using this technique, the percentage of isoprene‐emitting biomass varied from 20% to 42% (mean=31%; SD=8%). Responses of isoprene emission to varying light and temperature, measured on a sun‐adapted leaf of mango (Mangifera indica L.), suggest that existing algorithms developed for temperate species are adequate for tropical species as well. Incorporating these algorithms, estimates of isoprene‐emitting biomass, isoprene emission capacity, and site foliar biomass into a canopy flux model, canopy‐scale fluxes of isoprene were predicted and compared with the above‐canopy fluxes measured at two sites. Our bottom‐up approach overestimates fluxes by about 50%, but variations in measured fluxes between the two sites are largely explained by observed variation in the amount of isoprene‐emitting biomass.     

Leaf dynamics and insect herbivory in a Eucalyptus camaldulensis forest under moisture stress

Abstract The influence of soil moisture content on leaf dynamics and insect herbivory was examined between September 1991 and March 1992 in a river red gum (Eucalyptus camaldulensis) forest in southern central New South Wales. Long-term observations of leaves were made in trees standing either within intermittently flooded waterways or at an average of 37. 5m from the edge of the waterways. The mean soil moisture content was significantly (P≤0.05) greater in the waterways than in the non-flooded areas. Trees in the higher soil moisture regime produced significantly larger basal area increments and increased canopy leaf area. This increase in canopy leaf area was achieved, in part, through a significant increase in leaf longevity and mean leaf size. Although a greater number of leaves was initiated and abscissed per shoot from the non-flooded trees, more leaves were collected from litter traps beneath the denser canopies of the flooded trees. Consumption of foliage by insects on the trees subjected to flooding compared to the non-flooded trees was not significantly different. However, the relative impact of insect herbivory was significantly greater on the non-flooded trees. Leaf chewing was the most common form of damage by insects, particularly Chryso-melidae and Curculionidae. No species was present in outbreak during this study. Leaf survival decreased as the per cent area eaten per leaf increased. In addition, irrespective of the level of herbivory, leaf abscission tended to be higher in E. camaldulensis under moisture deficit. The influence of soil moisture content on the balance between river red gum growth and insect herbivory is discussed.     

Insect herbivory on mangrove leaves in North Queensland*

Estimates of leaf damage by insect herbivores are presented for 25 species of mangrove plants, comprising canopy and understorey species. Leaf area loss was highly variable among the species sampled, with means ranging from 0.3 to 35.0% of expanded leaf area. There was also great variability amongst leaves within species, and the mean coefficient of variation for leaf loss from the 25 species was 266%. Of the 12 species sampled at more than one site in North Queensland, eight exhibited small, but significant, between-site differences in herbivory. In general, it did not appear that height in the canopy influenced herbivory. For the dominant mangrove forest community type at Missionary Bay, an estimated mean of 2.1% of leaf production, or 11 g m^-2 per year, entered the direct grazing pathway. This very low figure is compared with estimates from other studies on mangrove forests and estimates from a variety of Australian terrestrial forests.     

A foliar disease model for use in wheat disease management decision support systems

A model of winter wheat foliar disease is described, parameterised and tested for Septoria tritici (leaf blotch), Puccinia striiformis (yellow rust), Erysiphe graminis (powdery mildew) and Puccinia triticina (brown rust). The model estimates disease‐induced green area loss, and can be coupled with a wheat canopy model, in order to estimate remaining light‐intercepting green tissue and hence the capacity for resource capture. The model differs from those reported by other workers in three respects. First, variables (such as weather, host resistance and inoculum pressure) that affect disease risk are integrated in their effect on disease progress. The agronomic and meteorological data called for are restricted to those commonly available to growers by their own observations and from meteorological service networks. Second, field observations during the growing season can be used both to correct current estimates of disease severity and to modify parameters that determine predicted severity. Third, pathogen growth and symptom expression are modelled to allow the effects of fungicides to be accounted for as protectant activity (reducing infections that occur postapplication) and eradicant activity (reducing growth of presymptomatic infections). The model was tested against data from a wide range of sites and varieties and was shown to predict the expected level of disease sufficiently accurately to support fungicide treatment decisions.     

Quantification of plant stress using remote sensing observations and crop models: the case of nitrogen management

Remote sensing techniques offer a unique solution for mapping stress and monitoring its time-course. This article reviews the main issues to be addressed for quantifying stress level from remote sensing observations, and to mitigate its impact on crop production by managing cultural practices. The case of nitrogen fertilization is used here as a paradigm. The derivation of canopy state variables such as the leaf area index (LAI) and chlorophyll content (C(ab)) is first addressed. It is demonstrated that the inversion of radiative transfer models leads to useful estimates of these variables. However, because of the ill-posed nature of the inverse problem, better accuracy is achieved when using prior information on the distribution of the variables and when multiplying LAI by C(ab) to get canopy level chlorophyll content. This variable, LAIxC(ab) is well suited for quantifying canopy level nitrogen content. It is used for nitrogen stress evaluation by comparison with a reference unstressed situation which is, however, not easy to get in practice. The combination of remote sensing observations with crop models provides an elegant solution for stress quantification through assimilation approaches. It fuses several sources of information within our knowledge of the processes involved and accounts for the environmental budget which can be integrated when making decisions about cultural practices. Conclusions are drawn on the issues related to the retrieval of canopy state variables from remote sensing data, to the link between these observables and crop models, and to the assimilation approaches. Avenues for further research are finally discussed along with the required observation system.     

Modelling canopy photosynthesis using parameters determined from simple non-destructive measurements

Most models for canopy photosynthesis require a large number of parameters as input which have to be determined by means of direct measurements. Such measurements are usually expensive, time consuming and destructive. The objective of the present study was, therefore, to develop a simple but accurate canopy photosynthesis model based on a minimum number of parameters that can be determined non-destructively. The results from previous studies were used to derive an empirical expression which describes the variation in leaf photosynthetic capacity (P_m) as a function of the light distribution in the canopy. The light distribution itself was calculated with a simple model which assumes only three leaf angle classes (0–30°, 30–60° and 60–90°). The leaf area index was determined indirectly from measurements of direct radiation below the canopy. The result was a model for canopy photosynthesis that requires only a few parameters. These parameters are the leaf photosynthetic capacity at the top of the canopy, the relative frequency of leaves in each of the three leaf angle classes, and the fraction of direct radiation below the canopy. Each of these parameters can be determined by means of simple non-destructive measurements. The model was applied to dense stands of two monocotyledonous species: rice (Oryza sativa L.) and pearl millet (Pennisetum americanum (L.) K. Schum.). The rates of canopy photosynthesis thus calculated were compared to those obtained with a more elaborate reference model. The differences between the values obtained with the two models were small. The present photosynthesis model can, therefore, be considered to be a suitable alternative for the more elaborate model. It was further discussed that, since the model is based on purely non-destructive measurements, it will be particularly useful in cases where it is required to estimate canopy photosynthesis at regular intervals over a length of time or in stands of vegetation that cover large areas of land.