J. Vos

Functional–structural plant modelling: a new versatile tool in crop science

Plants react to their environment and to management interventions by adjusting physiological functions and structure. Functional-structural plant models (FSPM), combine the representation of three-dimensional (3D) plant structure with selected physiological functions. An FSPM consists of an architectural part (plant structure) and a process part (plant functioning). The first deals with (i) the types of organs that are initiated and the way these are connected (topology), (ii) co-ordination in organ expansion dynamics, and (iii) geometrical variables (e.g. leaf angles, leaf curvature). The process part may include any physiological or physical process that affects plant growth and development (e.g. photosynthesis, carbon allocation). This paper addresses the following questions: (i) how are FSPM constructed, and (ii) for what purposes are they useful? Static, architectural models are distinguished from dynamic models. Static models are useful in order to study the significance of plant structure, such as light distribution in the canopy, gas exchange, remote sensing, pesticide spraying studies, and interactions between plants and biotic agents. Dynamic models serve quantitatively to integrate knowledge on plant functions and morphology as modulated by environment. Applications are in the domain of plant sciences, for example the study of plant plasticity as related to changes in the red:far red ratio of light in the canopy. With increasing availability of genetic information, FSPM will play a role in the assessment of the significance towards plant performance of variation in genetic traits across environments. In many crops, growers actively manipulate plant structure. FSPM is a promising tool to explore divergent management strategies.

Functional-Structural Plant Modelling in Crop Production

Functional-structural plant models (FSPMs) describe in quantitative terms the development over time of the three-dimensional (3D) structure of plants as governed by physiological processes and affected by environmental factors. FSPMs are particularly suited to analyse problems in which the spatial structure of the plant or its canopy is an essential factor to explain, e.g., plant competition (intra-plant, inter-plant, inter-species) and the effects of plant configuration and plant manipulation (e.g., pruning and harvesting) on yield and produce quality. This book describes the philosophy of functional-structural plant modelling and several tools for making FSPMs; it outlines methods for measuring essential parameters, including those pertaining to plant structure. As FSPMs offer new opportunities to model sinksource interactions, the physiological theory and modelling approaches regarding partitioning of carbon are given specific attention. Examples of application of FSPMs include wheat modelling in the context of remote sensing and the analysis of predatorprey insect interactions on glasshouse plants. The book will be useful for scientists and advanced students interested in innovative approaches in plant and crop modelling.

Using combined measurements of gas exchange and chlorophyll fluorescence to estimate parameters of a biochemical C3 photosynthesis model: a critical appraisal and a new integrated approach applied to leaves in a wheat (Triticum aestivum) canopy

We appraised the literature and described an approach to estimate the parameters of the Farquhar, von Caemmerer and Berry model using measured CO(2) assimilation rate (A) and photosystem II (PSII) electron transport efficiency (Phi(2)). The approach uses curve fitting to data of A and Phi(2) at various levels of incident irradiance (I(inc)), intercellular CO(2) (C(i)) and O(2). Estimated parameters include day respiration (R(d)), conversion efficiency of I(inc) into linear electron transport of PSII under limiting light [kappa(2(LL))], electron transport capacity (J(max)), curvature factor (theta) for the non-rectangular hyperbolic response of electron flux to I(inc), ribulose 1.5-bisphosphate carboxylase/oxygenase (Rubisco) CO(2)/O(2) specificity (S(c/o)), Rubisco carboxylation capacity (V(cmax)), rate of triose phosphate utilization (T(p)) and mesophyll conductance (g(m)). The method is used to analyse combined gas exchange and chlorophyll fluorescence measurements on leaves of various ages and positions in wheat plants grown at two nitrogen levels. Estimated S(c/o) (25 degrees C) was 3.13 mbar microbar(-1); R(d) was lower than respiration in the dark; J(max) was lower and theta was higher at 2% than at 21% O(2); kappa(2(LL)), V(cmax), J(max) and T(p) correlated to leaf nitrogen content; and g(m) decreased with increasing C(i) and with decreasing I(inc). Based on the parameter estimates, we surmised that there was some alternative electron transport.

The Plant TPX2 Protein Regulates Prospindle Assembly before Nuclear Envelope Breakdown

The Targeting Protein for Xklp2 (TPX2) is a central regulator of spindle assembly in vertebrate cells. The absence or excess of TPX2 inhibits spindle formation. We have defined a TPX2 signature motif that is present once in vertebrate sequences but twice in plants. Plant TPX2 is predominantly nuclear during interphase and is actively exported before nuclear envelope breakdown to initiate prospindle assembly. It localizes to the spindle microtubules but not to the interdigitating polar microtubules during anaphase or to the phragmoplast as it is rapidly degraded during telophase. We characterized the Arabidopsis thaliana TPX2-targeting domains and show that the protein is able to rescue microtubule assembly in TPX2-depleted Xenopus laevis egg extracts. Injection of antibodies to TPX2 into living plant cells inhibits the onset of mitosis. These results demonstrate that plant TPX2 already functions before nuclear envelope breakdown. Thus, plants have adapted nuclear–cytoplasmic shuttling of TPX2 to maintain proper spindle assembly without centrosomes.

Simulation of wheat growth and development based on organ-level photosynthesis and assimilate allocation

Intimate relationships exist between form and function of plants, determining many processes governing their growth and development. However, in most crop simulation models that have been created to simulate plant growth and, for example, predict biomass production, plant structure has been neglected. In this study, a detailed simulation model of growth and development of spring wheat (Triticum aestivum) is presented, which integrates degree of tillering and canopy architecture with organ-level light interception, photosynthesis, and dry-matter partitioning. An existing spatially explicit 3D architectural model of wheat development was extended with routines for organ-level microclimate, photosynthesis, assimilate distribution within the plant structure according to organ demands, and organ growth and development. Outgrowth of tiller buds was made dependent on the ratio between assimilate supply and demand of the plants. Organ-level photosynthesis, biomass production, and bud outgrowth were simulated satisfactorily. However, to improve crop simulation results more efforts are needed mechanistically to model other major plant physiological processes such as nitrogen uptake and distribution, tiller death, and leaf senescence. Nevertheless, the work presented here is a significant step forwards towards a mechanistic functional-structural plant model, which integrates plant architecture with key plant processes.

Understanding shoot branching by modelling form and function

Shoot branching plays a pivotal role in the development of the aboveground plant structure. Therefore, to understand branching in relation to the environment, it is not only necessary to integrate the knowledge on mechanisms that regulate branching at multiple levels of biological organisation, but also to include plant structure explicitly. To this end, we propose the application of an established methodology called functional-structural plant modelling.

Nutrient cycling in a cropping system with potato, spring wheat, sugar beet, oats and nitrogen catch crops. II. Effect of catch crops on nitrate leaching in autumn and winter

The Nitrate Directive of the European Union (EU) forces agriculture to reduce nitrate emission. The current study addressed nitrate emission and nitrate-N concentrations in leachate from cropping systems with and without the cultivation of catch crops (winter rye: Secale cereale L. and forage rape: Brassica napus ssp. oleifera (Metzg.) Sinksk). For this purpose, ceramic suction cups were used, installed at 80 cm below the soil surface. Soil water samples were extracted at intervals of ca 14 days over the course of three leaching seasons (September – February) in 1992–1995 on sandy soil in a crop rotation comprising potato (Solanum tuberosum L.), spring wheat (Triticum aestivum L.), sugar beet (Beta vulgaris L.) and oats (Avena sativa L.). Nitrate-N concentration was determined in the soil water samples. In a selection of samples several cations and anions were determined in order to analyze which cations primarily leach in combination with nitrate. The water flux at 80 cm depth was calculated with the SWAP model. Nitrate-N loss per interval was obtained by multiplying the measured nitrate-N concentration and the calculated flux. Accumulation over the season yielded the total nitrate-N leaching and the seasonal flux-weighted nitrate-N concentration in leachate. Among the cases studied, the total leaching of nitrate-N ranged between 30 and 140 kg ha–1. Over the leaching season, the flux-weighted nitrate-N concentration ranged between 5 and 25 mg L–1. Without catch crop cultivation, that concentration exceeded the EU nitrate-N standard (11.3 mg L–1) in all cases. Averaged for the current rotation, cultivation of catch crops would result in average nitrate-N concentrations in leachate near or below the EU nitrate standard. Nitrate-N concentrations correlated with calcium concentration and to a lesser extent with magnesium and potassium, indicating that these three ion species primarily leach in combination with nitrate. It is concluded that systematic inclusion of catch crops helps to decrease the nitrate-N concentration in leachate to values near or below the EU standard in arable rotations on sandy soils.

Phyllochron responds to acclimation to temperature and irradiance in maize

Crop models need accurate simulation of leaf canopy development. The thermal interval for leaf tip appearance (phyllochron) is critical for predicting the duration of vegetative development. The phyllochron in maize is shorter in temperate than in tropical and subtropical environments. As existing data has been evaluated in a narrow range of environments, the underlying mechanisms that affect phyllochron have not been adequately examined. The objectives of this study were to quantify the response of phyllochron to environmental variables and determine its stability across maize cultivars, to aid modelers in developing tools which accurately predict phenology. Maize was grown in field experiments at Wageningen, The Netherlands, Temple, Texas, USA, and three sites in Mexico, and in controlled environments at Wageningen. The experiment at Temple included grain sorghum and shading treatments to alter irradiance of the crop. Detailed data on leaf production and environmental conditions were collected. These data were combined with published data from field studies. Maize phyllochron acclimated to temperature and increased as mean daily temperature before tassel initiation increased from 12.5 to 25.5 degrees C, and increased in maize and sorghum in response to low irradiance. Temperature was the dominant influence, with phyllochron increasing by 1.7 degrees Cd per degrees C increase in daily mean temperature, as daily mean temperature before tassel initiation increased from 12.5 to 25.5 degrees C, and declined or remained constant when mean daily temperature before tassel initiation exceeded 25.5 degrees C. Only small differences in phyllochron occurred among cultivars. Phyllochron increased by 2 to 4 degrees Cd per MJ photosynthetically active radiation (PAR) as irradiance decreased from 9.6 to 1.1 MJ PAR m-2 day-1.

The contribution of phenotypic plasticity to complementary light capture in plant mixtures

Interspecific differences in functional traits are a key factor for explaining the positive diversity-productivity relationship in plant communities. However, the role of intraspecific variation attributable to phenotypic plasticity in diversity-productivity relationships has largely been overlooked. By taking a wheat (Triticum aestivum)-maize (Zea mays) intercrop as an elementary example of mixed vegetation, we show that plasticity in plant traits is an important factor contributing to complementary light capture in species mixtures. We conceptually separated net biodiversity effect into the effect attributable to interspecific trait differences and species distribution (community structure effect), and the effect attributable to phenotypic plasticity. Using a novel plant architectural modelling approach, whole-vegetation light capture was simulated for scenarios with and without plasticity based on empirical plant trait data. Light capture was 23% higher in the intercrop with plasticity than the expected value from monocultures, of which 36% was attributable to community structure and 64% was attributable to plasticity. For wheat, plasticity in tillering was the main reason for increased light capture, whereas for intercropped maize, plasticity induced a major reduction in light capture. The results illustrate the potential of plasticity for enhancing resource acquisition in mixed stands, and indicate the importance of plasticity in the performance of species-diverse plant communities.

Plant development and leaf area production in contrasting cultivars of maize grown in a cool temperate environment in the field

Crop models need accurate simulation of the interdependent processes of crop development and leaf area production. Crop development proceeds according to genotype characteristics and environmental influences, specifically temperature and photoperiod. It can be partly described by thermal requirements for development intervals and coefficients that describe genotype adaptation. The objectives of this study were to (a) quantify (i) time of tassel initiation, tasselling and silking; (ii) thermal intervals for initiation, appearance and expansion of successive leaves (iii) thermal duration from initiation to tip appearance and from tip appearance to collar appearance, and (iv) leaf area and canopy cover as measured by leaf area index (LAI) in contrasting cultivars of maize grown in the field in a cool environment; and (b) relate these to plant characteristics and environmental variables, particularly temperature. For these purposes, three cultivars of maize were grown in three and four cultivars in two serial plantings from 18 April to 24 June in field experiments at Wageningen, The Netherlands, in 1997, and detailed data on crop development, leaf production and environmental variables were collected. The base temperature (Tb) for maize was confirmed as 8 degrees C, but thermal time calculation needs to be re-examined to explore a recovery period after chilling injury. Equations that relate foliar properties to total leaf number and ordinal leaf position were derived. Individual leaf area can be described by the modified bell curve, and differences in temporal increase in LAI were related to parameters of leaf initiation, appearance and expansion.