A dynamical model of environmental effects on allocation to carbon-based secondary compounds in juvenile trees.

BACKGROUND AND AIMS: Patterns and variations in concentration of carbon-based secondary compounds in plant tissues have been explained by means of different complementary and, in some cases, contradictory plant defence hypotheses for more than 20 years. These hypotheses are conceptual models which consider environmental impacts on plant internal demands. In the present study, a mathematical model is presented, which converts and integrates the concepts of the 'Growth-Differentiation Balance' hypothesis and the 'Protein Competition' model into a dynamic plant growth model, that was tested with concentration data of polyphenols in leaves of juvenile apple, beech and spruce trees. The modelling approach is part of the plant growth model PLATHO that considers simultaneously different environmental impacts on the most important physiological processes of plants. METHODS: The modelling approach for plant internal resource allocation is based on a priority scheme assuming that growth processes have priority over allocation to secondary compounds and that growth-related metabolism is more strongly affected by nitrogen deficiency than defence-related secondary metabolism. KEY RESULTS: It is shown that the model can reproduce the effect of nitrogen fertilization on allocation patterns in apple trees and the effects of elevated CO(2) and competition in juvenile beech and spruce trees. The analysis of model behaviour reveals that large fluctuations in plant internal availability of carbon and nitrogen are possible within a single vegetation period. Furthermore, the model displays a non-linear allocation behaviour to carbon-based secondary compounds. CONCLUSIONS: The simulation results corroborate the underlying assumptions of the presented modelling approach for resource partitioning between growth-related primary metabolism and defence-related secondary metabolism. Thus, the dynamical modelling approach, which considers variable source and sink strengths of plant internal resources within different phenological growth stages, presents a successful translation of existing concepts into a dynamic mathematical model.

Analysis of competition effects in mono- and mixed cultures of juvenile beech and spruce by means of the plant growth simulation model PLATHO.

Inter- and intra-specific competition between plants for external resources is a critical process for plant growth in natural and managed ecosystems. We present a new approach to simulate competition for the resources light, water, and nitrogen between individual plants within a canopy. This approach was incorporated in a process-oriented plant growth simulation model. The concept of modelling competition is based on competition coefficients calculated from the overlap of occupied crown and soil volumes of each plant individual with the occupied volumes of its four nearest neighbours. The model was parameterised with data from a two-year phytotron experiment with juvenile beech and spruce trees growing in mono- and mixed cultures. For testing the model, an independent data set from this experiment and data from a second phytotron experiment with mixed cultures were used. The model was applied to analyse the consequences of start conditions and plant density on plant-plant competition. In both experiments, spruce dominated beech in mixed cultures. Based on model simulations, we postulate a large influence of start conditions and stand density on the outcome of the competition between the species. When both species have similar heights at the time of canopy closure, the model suggests a greater morphological plasticity of beech compared with spruce to be the crucial mechanism for competitiveness in mixed canopies. Similar to the experiment, in the model greater plasticity was a disadvantage for beech leading to it being outcompeted by the more persistent spruce.

The plant's capacity in regulating resource demand.

Regulation of resource allocation in plants is the key to integrate understanding of metabolism and resource flux across the whole plant. The challenge is to understand trade-offs as plants balance allocation between different and conflicting demands, e.g., for staying competitive with neighbours and ensuring defence against parasites. Related hypothesis evaluation can, however, produce equivocal results. Overcoming deficits in understanding underlying mechanisms is achieved through integrated experimentation and modelling the various spatio-temporal scaling levels, from genetic control and cell metabolism towards resource flux at the stand level. An integrated, interdisciplinary research concept on herbaceous and woody plants and its outcome to date are used, while drawing attention to currently available knowledge. This assessment is based on resource allocation as driven through plant-pathogen and plant-mycorrhizosphere interaction, as well as competition with neighbouring plants in stands, conceiving such biotic interactions as a "unity" in the control of allocation. Biotic interaction may diminish or foster effects of abiotic stress on allocation, as changes in allocation do not necessarily result from metabolic re-adjustment but may obey allometric rules during ontogeny. Focus is required on host-pathogen interaction under variable resource supply and disturbance, including effects of competition and mycorrhization. Cost/benefit relationships in balancing resource investments versus gains turned out to be fundamental in quantifying competitiveness when related to the space, which is subject to competitive resource exploitation. A space-related view of defence as a form of prevention of decline in competitiveness may promote conversion of resource turnover across the different kinds of biotic interaction, given their capacity in jointly controlling whole plant resource allocation.