A system-dynamic model on the competitive growth between Potamogeton malaianus Miq. and Spirogyra sp.

A system-dynamic model has been built to evaluate the competition between submerged macrophytes Potamogeton malaianus Miq. (PM) and filamentous green algae Spirogyra sp. (SP). The data background is based on a spring–summer and an autumn–winter experiment carried out in artificial field ponds. The experiments had the aim to acquire a knowledge base necessary to a successful restoration of submerged macrophyte vegetation in Lake Taihu, China by use of P. malaianus Miq. The model mainly focuses on variations in water volume; biomass dynamics of P. malaianus Miq., Spirogyra sp. and zoobenthos; nutrients cycling between water column, P. malaianus Miq., Spirogyra sp., zoobenthos, detritus and sediment. Sixteen state variables are included in the model: biomass of P. malaianus Miq., Spirogyra sp. and zoobenthos; nitrogen in sediments, detritus, in P. malaianus Miq., in Spirogyra sp. and zoobenthos; total dissolved nitrogen; phosphorus in sediments, detritus, in P. malaianus Miq., in Spirogyra sp. and in zoobenthos; total dissolved phosphorus, and water volume of the experiment pond. The calibration and validation of the model show a good accordance with the results of the spring–summer experiment and the autumn–winter experiment.     

Application of a Random Forest algorithm to predict spatial distribution of the potential yield of Ruditapes philippinarum in the Venice lagoon, Italy

We present a modelling framework that combines machine learning techniques and Geographic Information Systems to support the management of an important aquaculture species, Manila clam (Ruditapes philippinarum). We use the Venice lagoon (Italy), the first site in Europe for the production of R. philippinarum, to illustrate the potential of this modelling approach. To investigate the relationship between the yield of R. philippinarum and a set of environmental factors, we used a Random Forest (RF) algorithm. The RF model was tuned with a large data set (n = 1698) and validated by an independent data set (n = 841). Overall, the model provided good predictions of site-specific yields and the analysis of marginal effect of predictors showed substantial agreement among the modelled responses and available ecological knowledge for R. philippinarum. The most influent environmental factors for yield estimation were percentage of sand in the sediment, salinity, and water depth. Our results agree with findings from other North Adriatic lagoons. The application of the fitted RF model to continuous maps of all the environmental variables allowed estimates of the potential yield for the whole basin. Such a spatial representation enabled site-specific estimates of yield in different farming areas within the lagoon. We present a possible management application of our model by estimating the potential yield under the current farming distribution and comparing it to a proposed re-organization of the farming areas. Our analysis suggests a reduction of total yield is likely to result from the proposed re-organization.     

Modelling habitat overlap among sympatric mesocarnivores in southern Illinois,USA

Few researchers have developed large-scale habitat models for sympatric carnivore species. We created habitat models for red foxes (Vulpes vulpes), coyotes (Canis latrans) and bobcats (Lynx rufus) in southern Illinois, USA, using the Penrose distance statistic, remotely sensed landscape data, and sighting location data within a GIS. Our objectives were to quantify and spatially model potential habitat differences among species. Habitat variables were quantified for 1-km² buffered areas around mesocarnivore sighting locations. Following variable reduction procedures, five habitat variables (percentage of grassland patches, interspersion–juxtaposition of forest patches, mean fractal dimension of wetland patches and the landscape, and road density) were used for analysis. Only one variable differed (P < 0.05) between red fox and coyote sighting areas (road density) and bobcat and coyote sighting areas (mean fractal dimension of the landscape). However, all five variables differed between red fox and bobcat sighting areas, indicating considerable differences in habitat affiliation between this pair-group. Compared to bobcats, red fox sightings were affiliated with more grassland cover and larger grassland patches, higher road densities, lower interspersion and juxtaposition of forest patches, and lower mean fractal dimension of wetland patches. These differences can be explained by different life history requirements relative to specific cover types. We then used the Penrose distance statistic to create habitat models for red foxes and bobcats, respectively, based on the five-variable dataset. An independent set of sighting locations were used to validate these models; model fit was good with 65% of mesocarnivore locations within the top 50% of Penrose distance values. In general, red foxes were affiliated with mixtures of agricultural and grassland cover, whereas bobcats were associated with a combination of grassland, wetland, and forest cover. The greatest habitat overlap between red foxes and bobcats was found at the interface between forested areas and more open cover types. Our study provides insight into habitat overlap among sympatric mesocarnivores, and the distance-based modelling approach we used has numerous applications for modelling wildlife–habitat relationships over large scales.     

Imposed and inherent scales in cellular automata models of habitat

Both observational and modelling studies of the natural environment are characterised by their ‘grain’ and ‘extent’, the smallest and largest scales represented in time and space. These are imposed scales that should be chosen to ensure that the natural scales of the system are captured in the study. A simple cellular automata model of habitat represents only the presence or absence of vegetation, with global and local interactions described by four empirical parameters. Such a model can be formulated as a nonlinear Markov equation for the habitat probability. The equation produces inherent space and time scales that may be considered as transition scales or the scales for recovery from disturbance. However, if the resolution of the model is changed, the empirical parameters must be changed to preserve the properties of the system. Further, changes in the spatial resolution lead to different interpretations of the spatial structure. In particular, as the resolution is reduced, the apparent dominance of one habitat type over the other increases. The model provides an ability to compare both field and model investigations conducted at different resolutions in time and space.     

Evaluating weather effects on interannual variation in net ecosystem productivity of a coastal temperate forest landscape: A model intercomparison

Forest productivity is strongly affected by seasonal weather patterns and by natural or anthropogenic disturbances. However weather effects on forest productivity are not currently represented in inventory-based models such as CBM-CFS3 used in national forest C accounting programs. To evaluate different approaches to modelling these effects, a model intercomparison was conducted among CBM-CFS3 and four process models (ecosys, CN-CLASS, Can-IBIS and 3PG) over a 2500 ha landscape in the Oyster River (OR) area of British Columbia, Canada. The process models used local weather data to simulate net primary productivity (NPP), net ecosystem productivity (NEP) and net biome productivity (NBP) from 1920 to 2005. Other inputs used by the process and inventory models were generated from soil, land cover and disturbance records. During a period of intense disturbance from 1928 to 1943, simulated NBP diverged considerably among the models. This divergence was attributed to differences among models in the sizes of detrital and humus C stocks in different soil layers to which a uniform set of soil C transformation coefficients was applied during disturbances. After the disturbance period, divergence in modelled NBP among models was much smaller, and attributed mainly to differences in simulated NPP caused by different approaches to modelling weather effects on productivity. In spite of these differences, age-detrended variation in annual NPP and NEP of closed canopy forest stands was negatively correlated with mean daily maximum air temperature during July-September (T_amax) in all process models (R² = 0.4-0.6), indicating that these correlations were robust. The negative correlation between T_amax and NEP was attributed to different processes in different models, which were tested by comparing CO₂ fluxes from these models with those measured by eddy covariance (EC) under contrasting air temperatures (T_a). The general agreement in sensitivity of annual NPP to T_amax among the process models led to the development of a generalized algorithm for weather effects on NPP of coastal temperate coniferous forests for use in inventory-based models such as CBM-CFS3: NPP′ = NPP − 57.1 (T_amax − 18.6), where NPP and NPP′ are the current and temperature-adjusted annual NPP estimates from the inventory-based model, 18.6 is the long-term mean daily maximum air temperature during July-September, and T_amax is the mean value for the current year. Our analysis indicated that the sensitivity of NPP to T_amax was nonlinear, so that this algorithm should not be extrapolated beyond the conditions of this study. However the process-based methodology to estimate weather effects on NPP and NEP developed in this study is widely applicable to other forest types and may be adopted for other inventory based forest carbon cycle models.     

SPAnDX: a process-based population dynamics model to explore management and climate change impacts on an invasive alien plant,Acacia nilotica

This paper describes a process-based metapopulation dynamics and phenology model of prickly acacia, Acacia nilotica, an invasive alien species in Australia. The model, SPAnDX, describes the interactions between riparian and upland sub-populations of A. nilotica within livestock paddocks, including the effects of extrinsic factors such as temperature, soil moisture availability and atmospheric concentrations of carbon dioxide. The model includes the effects of management events such as changing the livestock species or stocking rate, applying fire, and herbicide application. The predicted population behaviour of A. nilotica was sensitive to climate. Using 35 years daily weather datasets for five representative sites spanning the range of conditions that A. nilotica is found in Australia, the model predicted biomass levels that closely accord with expected values at each site. SPAnDX can be used as a decision-support tool in integrated weed management, and to explore the sensitivity of cultural management practices to climate change throughout the range of A. nilotica. The cohort-based DYMEX modelling package used to build and run SPAnDX provided several advantages over more traditional population modelling approaches (e.g. an appropriate specific formalism (discrete time, cohort-based, process-oriented), user-friendly graphical environment, extensible library of reusable components, and useful and flexible input/output support framework).     

Predicting reproduction rate of varroa

A single equation is derived to predict population-density effects on the reproduction rate of the honey bee parasite Varroa destructor Anderson and Trueman. This equation provides a simpler alternative to the approach currently used in the literature, and additionally corrects an anomaly in that approach. The method is then extended to the case of co-existing haplotypes of Varroa. It thus derives an equation used without proof for modelling biocontrol of Varroa, and examines the error caused by an approximation necessary for a closed form solution. Additionally a varroa population model incorporating the derived equation is described.     

Sanitary felling of Norway spruce due to spruce bark beetles in Slovenia: A model and projections for various climate change scenarios

A model is presented to predict sanitary felling of Norway spruce (Picea abies) due to spruce bark beetles (Ips typographus, Pityogenes chalcographus) in Slovenia according to different climate change scenarios. The model incorporates 21 variables that are directly or indirectly related to the dependent variable, and that can be arranged into five groups: climate, forest, landscape, topography, and soil. The soil properties are represented by 8 variables, 4 variables define the topography, 4 describe the climate, 4 define the landscape, and one additional variable provides the quantity of Norway spruce present in the model cell. The model was developed using the M5′ model tree. The basic spatial unit of the model is 1 km², and the time resolution is 1 year. The model evaluation was performed by three different measures: (1) the correlation coefficient (51.9%), (2) the Theil's inequality coefficient (0.49) and (3) the modelling efficiency (0.32). Validation of the model was carried out by 10-fold cross-validation. The model tree consists of 28 linear models, and model was calculated for three different climate change scenarios extending over a period until 2100, in 10-year intervals. The model is valid for the entire area of Slovenia; however, climate change projections were made only for the Maribor region (596 km²). The model assumes that relationships among the incorporated factors will remain unchanged under climate change, and the influence of humans was not taken into account. The structure of the model reveals the great importance of landscape variables, which proved to be positively correlated with the dependent variable. Variables that describe the water regime in the model cell were also highly correlated with the dependent variable, with evapotranspiration and parent material being of particular importance. The results of the model support the hypothesis that bark beetles do greater damage to Norway spruce artificially planted out of its native range in Slovenia, i.e., lowlands and soils rich in N, P, and K. The model calculation for climate change scenarios in the Maribor region shows an increase in sanitary felling of Norway spruce due to spruce bark beetles, for all scenarios. The model provides a path towards better understanding of the complex ecological interactions involved in bark beetle outbreaks. Potential application of the results in forest management and planning is discussed.     

Multi-metric evaluation of the models WARM,CropSyst, and WOFOST for rice

WARM (Water Accounting Rice Model) simulates paddy rice (Oryza sativa L.), based on temperature-driven development and radiation-driven crop growth. It also simulates: biomass partitioning, floodwater effect on temperature, spikelet sterility, floodwater and chemicals management, and soil hydrology. Biomass estimates from WARM were evaluated and compared with the ones from two generic crop models (CropSyst, WOFOST). The test-area was the Po Valley (Italy). Data collected at six sites from 1989 to 2004 from rice crops grown under flooded and non-limiting conditions were split into a calibration (to estimate some model parameters) and a validation set. For model evaluation, a fuzzy-logic based multiple-metrics indicator (MQI) was used: 0 (best) ≤ MQI ≤ 1 (worst). WARM estimates compared well with the actual data (mean MQI = 0.037 against 0.167 and 0.173 with CropSyst and WOFOST, respectively). On an average, the three models performed similarly for individual validation metrics such as modelling efficiency (EF > 0.90) and correlation coefficient (R > 0.98). WARM performed best in a weighed measure of the Akaike Information Criterion: (worst) 0<wk<1$0<w_k<1$ (best), considering estimation accuracy and number of parameters required to achieve it (mean wk=0.983$w_k=0.983$ against 0.007 and ∼0.000 with CropSyst and WOFOST, respectively). WARM results were sensitive to 30% of the model parameters (ratio being lower with both CropSyst, <10%, and WOFOST, <20%), but appeared the easiest model to use because of the lowest number of crop parameters required (10 against 15 and 34 with CropSyst and WOFOST, respectively). This study provides a concrete example of the possibilities offered using a range of assessment metrics to evaluate model estimates, predictive capabilities, and complexity.     

A multi-agent simulation to assess the risk of malaria re-emergence in southern France

A multi-agent simulation (MAS) was developed to assess the risk of malaria re-emergence in the Camargue in southern France, a non-endemic area where mosquitoes of the genus Anopheles (Culicidae) live. The contact rate between people and potential malaria vectors, or the human biting rate, is one of the key factor to predict the risk of re-emergence of malaria, would the parasite be introduced in the region. Our model (called MALCAM) represents the different agents that could influence malaria transmission in the Camargue – people, mosquitoes, animal hosts and the landscape – in a spatially explicit environment. The model simulates spatial and temporal variations in human biting rate at the landscape scale. These variations depend on the distribution of people and potential vectors, their behaviour and their interactions. A land use/cover map was used as a cellular-spatial support for the movements of and interactions between mobile agents. The model was tested for its sensitivity to variations in parameter values, and for the agreement between field observations and model predictions. The MALCAM model provides a tool to better understand the interactions between the multiple agents of the disease transmission system, and the land use and land cover factors that control the spatial heterogeneity in these interactions. It allows testing hypotheses and scenarios related to disease dynamics by varying the value of exogenous biological, geographical, or human factors. This application of agent-based modelling to a human vector-borne disease can be adapted to different diseases and regions.     

Modelling N mineralization from green manure and farmyard manure from a laboratory incubation study

Predicting N mineralization from organic manures like farmyard manure (FYM) is more difficult than from fresh organic materials like crop residues, as the manures vary greatly in composition. A laboratory incubation experiment was carried out for 98 days at 30 °C under aerobic conditions to study the effects on N dynamics of Gliricidia (Gliricidia sepium, Jacquin) and FYM application to soil at 5 and 10 g kg⁻¹. Application of Gliricidia induced N mineralization from the start of incubation period, with the amount of N mineralized increasing with rate of application. In contrast, application of FYM resulted in immobilization of mineral N in soil, irrespective of the rate of application. The initial net immobilization from FYM was limited by availability of N in the soil for the higher rate of application.We used the APSIM SoilN module to simulate these contrasting patterns of mineralization of N from Gliricidia and from FYM. The prediction of N mineralized from Gliricidia was better than FYM. The default model parameters specify that the fresh organic matter pools (FPOOL1, FPOOL2 and FPOOL3) have the same C:N ratio and this assumption was ineffective in predicting N mineralized from FYM. The predictive ability of the model improved when this default assumption was modified based on the size of the individual pools (FPOOL1, FPOOL2 and FPOOL3), and the pool's C:N ratios. The modelling efficiency, a measure of goodness of fit between the simulated and observed data, improved markedly for the modified model. The discrepancy between the modelled and observed data was a tendency for the model to underestimate the rate of re-mineralization at the lower rate of application of FYM in the later part of incubation. Unfortunately the appropriate modification to the size and C:N ratios of the FPOOLs could not be determined on the basis of chemical analysis alone. Thus, a true predictive application of the model to a new FYM material is not yet possible.     

Ecological relevance of performance criteria for species distribution models

Species distribution models have often been developed based on ecological data. To develop reliable data-driven models, however, a sound model training and evaluation procedures are needed. A crucial step in these procedures is the assessment of the model performance, with as key component the applied performance criterion. Therefore, we reviewed seven performance criteria commonly applied in presence-absence modelling (the correctly classified instances, Kappa, sensitivity, specificity, the normalised mutual information statistic, the true skill statistic and the odds ratio) and analysed their application in both the model training and evaluation process. Although estimates of predictive performance have been used widely to assess final model quality, a systematic overview was missing because most analyses of performance criteria have been empirical and only focused on specific aspects of the performance criteria. This paper provides such an overview showing that different performance criteria evaluate a model differently and that this difference may be explained by the dependency of these criteria on the prevalence of the validation set. We showed theoretically that these prevalence effects only occur if the data are inseparable by an n-dimensional hyperplane, n being the number of input variables. Given this inseparability, different performance criteria focus on different aspects of model performance during model training, such as sensitivity, specificity or predictive accuracy. These findings have important consequences for ecological modelling because ecological data are mostly inseparable due to data noise and the complexity of the studied system. Consequently, it should be very clear which aspect of the model performance is evaluated, and models should be evaluated consistently, that is, independent of, or taking into account, species prevalence. The practical implications of these findings are clear. They provide further insight into the evaluation of ecological presence/absence models and attempt to assist modellers in their choice of suitable performance criteria.     

Application of a temperature-dependent von Bertalanffy growth model to bullhead (Cottus gobio)

The most studied and commonly applied model of fish growth is the von Bertalanffy model. However, this model does not take water temperature into account, which is one of the most important environmental factors affecting the life cycle of fish, as many physiological processes that determine growth, e.g. metabolic rate and oxygen supply, are directly influenced by temperature. In the present study we propose a version of the von Bertalanffy growth model that includes mean annual water temperatures by correlating the growth coefficient, k, explicitly and the asymptotic length, L_∞, implicitly to water temperature. All relationships include parameters with an obvious biological relevance that makes them easier to identify. The model is used to fit growth data of bullhead (Cottus gobio) at different locations in the Bez River network (Drme, France). We show that temperature explains much of the growth variability at the different sampling sites of the network.     

Theoretical analysis of change in forest carbon use efficiency with stand development: A case study on Hinoki Cypress (Chamaecyparis obtusa (Sieb. et Zucc.) Endl.) plantation from the seedling stage

Changes in carbon use efficiency (CUE), which is defined as the ratio of net primary production (NPP) to gross primary production (GPP), were estimated for the aerial parts of the Hinoki Cypress (Chamaecyparis obtusa (Sieb. et Zucc.) Endl.) with respect to stand development. The analysis incorporated previously published data from the early stages of stand development, namely the seedling stages of the cypress. For this analysis, a simple mathematical model to assess the changes in CUE was developed by incorporating data on physiological variables and mass of woody species. The CUE tended to increase with increases in the aboveground biomass of the stand, and then decreased gradually despite increases in the aboveground biomass. The CUE-value (0.28, 0.39) of the seedling stage was lower than that (0.33-0.58) of the young or mature trees. To examine the effect of physiological variables and mass on CUE, the ratios of the specific respiration rate to the specific photosynthetic rate (r/a) and the leaf biomass to the aboveground biomass or leaf mass ratio (y_L/y_T) were calculated. The low value of CUE at the seedling stage was due to the high ratio of specific respiration rate to specific photosynthetic rate r/a, but was not due to the high value of the leaf mass ratio y_L/y_T. In addition, the decline in CUE associated with older stages of stand development was due to the decreasing changes in y_L/y_T, and the r/a ratio did not influence the change in CUE.     

Development of a hydrothermal time seed germination model which uses the Weibull distribution to describe base water potential

Seed germination has been modelled extensively using hydrothermal time (HTT) models, that predict time to germination as a function of the extent to which seedbed temperature, T, and water potential, Ψ, exceed the base temperature, T_b, and base water potential, Ψ_b, of each seed percentile, g. Within a seed population the variation in time to germination arises from variation in Ψ_b(g) modelled by a normal distribution. We tested the assumption of normality in the distribution of Ψ_b(g) by germinating seed of two unrelated species with non-dormant seed (Buddleja davidii (Franch.) and Pinus radiata D. Don) across a range of constant Ψ at sub-optimal T. When incorporated into a HTT model the Weibull distribution more accurately described both the right skewed distribution of Ψ_b(g) and germination time course over sub-optimal T than the HTT based on the normal distribution, for both species. Given the flexibility of the Weibull distribution this model not only provides a useful method for predicting germination but also a means of determining the distribution of Ψ_b(g).     

A process-based population dynamics model to explore target and non-target impacts of a biological control agent

The risks and benefits associated with efforts to control invasive alien species using classical biological control are being subjected to increasing scrutiny. A process-based population dynamics model was developed to explore the interactions between a folivorous biological control agent, Cleopus japonicus, and its plant host Buddleja davidii. The model revealed that climate could have a significant impact upon the interactions between B. davidii and C. japonicus. At the coolest sites, the impact of C. japonicus on B. davidii was slowed, but it was still eventually capable of controlling populations of B. davidii. At the warmer sites where both B. davidii and C. japonicus grew faster, B. davidii succumbed rapidly to weevil damage. We hypothesise that barring an encounter with a natural enemy, C. japonicus will eventually be able to provide sustained control B. davidii throughout the North Island of New Zealand. The model scenarios illustrate the potential for the C. japonicus population to attain high densities rapidly, and to defoliate patches of B. davidii, creating the potential for spill-over feeding on non-target plants. The potential magnitude of this threat will depend partly on the climate suitability for C. japonicus, the pattern by which it migrates in response to a reduction in the available leaf resource, and the suitability of non-target plants as hosts. In all migration scenarios considered, the pattern of population growth and resource consumption by C. japonicus was exponential, with a strong tendency toward complete utilisation of resource patches more quickly at the warmer compared to colder sites. In addition to providing some useful hypotheses about the effects of climate on the biological control system, and the non-target risks, it also provides some insight into the mechanisms by which climate affects the system.     

Computer simulation applied to the biological control of the insect Aphis gossypii for the parasitoid Lysiphlebus testaceipes

In integrated pest management (IPM), biological control is one of the possible options for the prevention or remediation of an unacceptable pest activity or damage. The success of forecast models in IPM depends, among other factors, on the knowledge of temperature effect over pests and its natural enemies. In this work, we simulated the effects of parasitism of Lysiphlebus testaceipes (Cresson, 1880) (Hymenoptera: Aphidiidae) on Aphis gossypii (Glover, 1877) (Hemiptera: Aphididae), a pest that is associated to crops of great economic importance in several parts of the world. We made use of experimental data relative to the host and its parasitoid at different temperatures. Age structure was incorporated into the dynamics through the Penna model. The results obtained showed that simulation, as a forecast model, can be a useful tool for biological control programs.     

Using geospatial business intelligence paradigm to design a multidimensional conceptual model for efficient coastal erosion risk assessment

One of the main challenges in Coastal Erosion Risk Assessment (CERA) is integrating and analysis of conflicting data in various time periods and spatial scales through dissimilar environmental, social, and economic criteria. Currently, Geographical Information Systems (GIS) are widely used in risk assessment despite their drawbacks and limitations as transactional systems for multi-scales, multi-epochs, and multi-themes analysis. Hence, an analytical conceptual framework is proposed in this paper based on geospatial business intelligence paradigm to develop a Spatial Multidimensional Conceptual Model (SMCM) to assess coastal erosion risk. The model is designed based on Spatial On-Line Analytical Processing (SOLAP) platform, on the top of both analytical and transactional paradigms, to allow fast synthesis of cross-tabulated data and easy comparisons over space, scales, epochs, and themes. This objective is achieved through a comprehensive integration of multiple environmental, social, and economic criteria as well as their interactions at various scales. It also takes into account multiple elements at risk such as people, infrastructure, and built environment as different dimensions of analysis. Using this solution allows decision makers to benefit from on-demand, interactive, and comprehensive information in a way that is not possible using GIS alone. The developed model can easily be adapted for any other coastal region through the proposed framework to perform risk assessment. The advantages and drawbacks of the proposed framework are also discussed and new research perspectives are presented.     

Integrating effects of leaf nitrogen,age, rank,and growth temperature into the photosynthesis-stomatal conductance model LEAFC3-N parameterised for barley (Hordeum vulgare L.)

A crucial challenge for including biophysical photosynthesis–transpiration models into complex crop growth models is to integrate the plasticity of photosynthetic processes that is related to factors like nitrogen (N) content, age, and rank of leaves, or to the adaptation of plants to growth temperature (T_g). Here we present a new version of the combined photosynthesis-stomatal conductance model LEAFC3-N [Müller, J., Wernecke, P., Diepenbrock, W., 2005. LEAFC3-N: a nitrogen sensitive extension of the CO₂ and H₂O gas exchange model LEAFC3 parameterised and tested for winter wheat (Triticum aestivum L.). Ecological Modelling 183, 183–210.] that was revised, extended and completely re-parameterised for barley (Hordeum vulgare L.) with special regard for these factors to facilitate the use of the model in ecophysiological studies and in crop modelling. The analysis is based on novel comprehensive data on photosynthetic CO₂ and light response curves measured at two oxygen concentrations and different temperatures on leaves of barley (H. vulgare L.) differing in leaf N and chlorophyll content. Plants were grown in climatic chambers or in the field at different N and T_g.We thoroughly revised the existing and introduced new nitrogen relations for key model parameters that account for a linear increase with leaf N of V_max, J_max, T_p, and R_dmax (maximum rates of carboxylation, electron transport, triose phosphate export, and mitochondrial respiration), a saturation-type increase of φ (quantum yield of electron transport), and a non-linear decrease of θ and m (curvature of the light dependence of electron transport rate, scaling factor of the stomata model). The adaptation of photosynthetic characteristics to T_g was included into the model by linear relations that were observed between T_g and the activation energy ΔH_a of the temperature response characteristics of V_max, J_max, and T_p as well as of the nitrogen dependency of these characteristics. Based on an analysis of diurnal time courses of gas exchange rates it was found necessary including not only the relation between leaf water potential (Ψ) and stomatal conductance as used originally in LEAFC3, but additional effects on V_max and J_max. With the above-listed extensions, the model was capable to reproduce the observed plasticity and the recorded diurnal time courses of gas exchange rates fairly well. Thus, we conclude that the new model version can be used under a broad range of conditions, both for ecophysiological studies and as a submodel of crop growth models. The results presented here for barley will facilitate adapting photosynthesis models like LEAFC3-N to other C₃-species as well. The modelling of the effects of drought stress should be further elaborated in future based on more specific experiments.     

Relationships between simulated water stress and mortality and growth rates in underplanted Toona ciliata Roem. in subtropical Argentinean plantations

Toona ciliata Roem. (Australian red cedar) requires a nurse-tree overstory to prevent damage from drought and irradiation in some regions of north-eastern Argentina. T. ciliata was planted in the understory of Pinus taeda L. (625 stems/ha), Pinus elliottii Engelm. × Pinus caribaea Morelet (625 stems/ha), and Grevillea robusta A. Cunn. (833 stems/ha) nurse trees, which were thinned to 0, 25, 50, 75 and 100% of the initial densities. We measured initial T. ciliata mortality and growth as well as Leaf Area Index (LAI) based on light transmission. T. ciliata soil water availability and its effect on early growth and mortality were examined by modelling drought stress using the two-dimensional forest hydrology model ForWaDy. Simulated patterns in T. ciliata water stress for the different overstory treatments were consistent with observed patterns of mortality. Early mortality was lowest with a G. robusta overstory, with corresponding lowest drought stress values and high modelled soil water contents in the top soil layer in intermediate and high overstory densities. Mortality was highest with a P. elliottii × P. caribaea overstory in treatments with the highest modelled drought stress values in the most open treatments. The model supported our field observations by indicating that water stress was an important limitation to T. ciliata survival and growth on our study sites. The linkage between T. ciliata establishment success, early growth and soil water availability as indicated by ForWaDy, leads us to conclude that the model is a suitable stand management tool for guiding establishment of T. ciliata plantations.