Integrating terrestrial laser scanning with functional-structural plant models to investigate ecological and evolutionary processes of forest communities.

BACKGROUND: Woody plants (trees and shrubs) play an important role in terrestrial ecosystems, but their size and longevity make them difficult subjects for traditional experiments. In the last 20 years functional-structural plant models (FSPMs) have evolved: they consider the interplay between plant modular structure, the immediate environment and internal functioning. However, computational constraints and data deficiency have long been limiting factors in a broader application of FSPMs, particularly at the scale of forest communities. Recently, terrestrial laser scanning (TLS), has emerged as an invaluable tool for capturing the 3-D structure of forest communities, thus opening up exciting opportunities to explore and predict forest dynamics with FSPMs. SCOPE: The potential synergies between TLS-derived data and FSPMs have yet to be fully explored. Here, we summarize recent developments in FSPM and TLS research, with a specific focus on woody plants. We then evaluate the emerging opportunities for applying FSPMs in an ecological and evolutionary context, in light of TLS-derived data, with particular consideration of the challenges posed by scaling up from individual trees to whole forests. Finally, we propose guidelines for incorporating TLS data into the FSPM workflow to encourage overlap of practice amongst researchers. CONCLUSIONS: We conclude that TLS is a feasible tool to help shift FSPMs from an individual-level modelling technique to a community-level one. The ability to scan multiple trees, of multiple species, in a short amount of time, is paramount to gathering the detailed structural information required for parameterizing FSPMs for forest communities. Conventional techniques, such as repeated manual forest surveys, have their limitations in explaining the driving mechanisms behind observed patterns in 3-D forest structure and dynamics. Therefore, other techniques are valuable to explore how forests might respond to environmental change. A robust synthesis between TLS and FSPMs provides the opportunity to virtually explore the spatial and temporal dynamics of forest communities.

Photosynthetical activity modelisation of olive trees growing under drought conditions.

Predicting photosynthetic production in olive trees is a key feature in managing the effect of climate change on arid areas. Functional-structural plant modelling is a promising tool for achieving this goal. We used a photosynthetic sub-model that accounted for water and temperature stress and implemented it into LIGNUM model. We then conducted an experiment to validate the model at the leaf level using olive trees (Olea europaea) grown under various climatic condition. Then, we simulated photosynthetic production of three static olive tree models aged 1, 2, and 3 years. Results revealed a good fit between observed and predicted photosynthesis, with coefficient of determination (R2) values of 0.94 and 0.93 for Chemlali and Zarrazi cultivars, respectively. These results showed that the impact of water stress on photosynthetic production was marginal.

A study of crown development mechanisms using a shoot-based tree model and segmented terrestrial laser scanning data.

Background and Aims: Functional-structural plant models (FSPMs) allow simulation of tree crown development as the sum of modular (e.g. shoot-level) responses triggered by the local environmental conditions. The actual process of space filling by the crowns can be studied. Although the FSPM simulations are at organ scale, the data for their validation have usually been at more aggregated levels (whole-crown or whole-tree). Measurements made by terrestrial laser scanning (TLS) that have been segmented into elementary units (internodes) offer a phenotyping tool to validate the FSPM predictions at levels comparable with their detail. We demonstrate the testing of different formulations of crown development of Scots pine trees in the LIGNUM model using segmented TLS data. Methods: We made TLS measurements from four sample trees growing in a forest on a relatively poor soil from sapling size to mature stage. The TLS data were segmented into internodes. The segmentation also produced information on whether needles were present in the internode. We applied different formulations of crown development (flushing of buds and length of growth of new internodes) in LIGNUM. We optimized the parameter values of each formulation using genetic algorithms to observe the best fit of LIGNUM simulations to the measured trees. The fitness function in the estimation combined both tree-level characteristics (e.g. tree height and crown length) and measures of crown shape (e.g. spatial distribution of needle area). Key Results: Comparison of different formulations against the data indicates that the Extended Borchert-Honda model for shoot elongation works best within LIGNUM. Control of growth by local density in the crown was important for all shoot elongation formulations. Modifying the number of lateral buds as a function of local density in the crown was the best way to accomplish density control. Conclusions: It was demonstrated how segmented TLS data can be used in the context of a shoot-based model to select model components.

Contributions of leaf photosynthetic capacity, leaf angle and self-shading to the maximization of net photosynthesis in Acer saccharum: a modelling assessment.

BACKGROUND AND AIMS: Plants are expected to maximize their net photosynthetic gains and efficiently use available resources, but the fundamental principles governing trade-offs in suites of traits related to resource-use optimization remain uncertain. This study investigated whether Acer saccharum (sugar maple) saplings could maximize their net photosynthetic gains through a combination of crown structure and foliar characteristics that let all leaves maximize their photosynthetic light-use efficiency (ε). METHODS: A functional-structural model, LIGNUM, was used to simulate individuals of different leaf area index (LAI(ind)) together with a genetic algorithm to find distributions of leaf angle (L(A)) and leaf photosynthetic capacity (A(max)) that maximized net carbon gain at the whole-plant level. Saplings grown in either the open or in a forest gap were simulated with A(max) either unconstrained or constrained to an upper value consistent with reported values for A(max) in A. saccharum. KEY RESULTS: It was found that total net photosynthetic gain was highest when whole-plant PPFD absorption and leaf ε were simultaneously maximized. Maximization of ε required simultaneous adjustments in L(A) and A(max) along gradients of PPFD in the plants. When A(max) was constrained to a maximum, plants growing in the open maximized their PPFD absorption but not ε because PPFD incident on leaves was higher than the PPFD at which ε(max) was attainable. Average leaf ε in constrained plants nonetheless improved with increasing LAI(ind) because of an increase in self-shading. CONCLUSIONS: It is concluded that there are selective pressures for plants to simultaneously maximize both PPFD absorption at the scale of the whole individual and ε at the scale of leaves, which requires a highly integrated response between L(A), A(max) and LAI(ind). The results also suggest that to maximize ε plants have evolved mechanisms that co-ordinate the L(A) and A(max) of individual leaves with PPFD availability.

Toward extension of a single tree functional-structural model of Scots pine to stand level: effect of the canopy of randomly distributed, identical trees on development of tree structure.

Functional-structural plant growth models (FSPMs) combine the description of the structure of plants and the resource acquisition and partitioning at a detailed architectural level. They offer a means to study tree and stand development on the basis of a structurally accurate description that combines resource capture at the same level of detail. We describe here how a 'shoot-based' individual tree model, LIGNUM of Scots pine (Pinus sylvestris L.) has been applied to a group of identical trees (forest). The model has been applied to isolated trees and saplings growing in forest gaps. First, we present the LIGNUM model and the changes necessary for simulation of a forest instead of individual trees. LIGNUM derives tree growth on the basis of a process-based model of tree carbon balance and the architectural development of the 3-D tree crown. The time step is 1 year. We realised the forest as consisting of individual Scots pine trees on a plot 17 × 17 m, but simplified the stand description by simulating the growth of only one tree in the middle of the plot and assumed that the other trees were identical to it at all times. The model produced results that are comparable with observations made in real Scots pine trees and tree stands in Finland. The simulations with variable values of the parameters controlling the foliage-sapwood relationship, amount of sapwood required below a point in a branch or a stem, and the senescence of sapwood showed how growth declines when the sapwood requirement in the branches and stem was high. In this case, the proportion of resources allocated to the needles became small and the needle mass was low.

Shoot growth and crown development: effect of crown position in three-dimensional simulations.

Trees have been increasingly considered as modular organisms, with individual shoots forming autonomous units that respond semi-independently to their surrounding environment. However, there is evidence for fairly strict hormonal control of tree crown development. Studies on the hydraulic architecture of trees suggest a closer functional connection between shoots and crown development than is postulated by the theory of branch autonomy. We studied how shoot growth pattern influences growth and crown architecture in young Scots pine trees simulated by the LIGNUM model assuming that (a) the growth of a shoot mainly depends on its light climate and (b) the growth of a shoot is influenced by its position within the crown. We determined shoot position within the crown based on a recently developed vigor index. The vigor index compares the relative axis cross-sectional area from the base of the tree to each shoot and gives a value of 1 to the pathway of the greatest cross-sectional area. All other shoots attain values between 0 and 1 depending on their cross-sectional areas and the cross-sectional areas of the branches leading there from the main axis. The shoot light climate is characterized by annually intercepted photosynthetically active radiation. We compared the results from simulations (a) and (b) against an independent data set. The addition of a within-shoot position index (the vigor index) to our simulation (simulation b) resulted in a more realistic tree form than that obtained with simulation (a) alone. We discuss the functional significance of the results as well as the possibilities of using an index of shoot position in simulations of crown architecture.