The community of root fungi is associated with the growth rate of Norway spruce (Picea abies).

Our study delved into the relationship between root-associated fungi, gene expression and plant morphology in Norway spruce cuttings derived from both slow-and fast-growing trees. We found no clear link between the gene expression patterns of adventitious roots and the growth phenotype, suggesting no fundamental differences in the receptiveness to fungal symbionts between the phenotypes. Interestingly, saplings from slow-growing parental trees exhibited a higher richness of ectomycorrhizal species and larger roots. Some ectomycorrhizal species, typically found on mature spruces, were more prevalent on saplings from slow-growing spruces. The ericoid mycorrhizal fungus, Hyaloscypha hepaticola, showed a stronger association with saplings from fast-growing spruces. Moreover, saplings from slow-growing spruces had a greater number of Ascomycete taxa and free-living saprotrophic fungi. Aboveground sapling stems displayed some phenotypic variation; saplings from fast-growing phenotypes had longer branches but fewer whorls in their stems compared to those from the slow-growing group. In conclusion, the observed root-associated fungi and phenotypic characteristics in young Norway spruces may play a role in their long-term growth rate. This suggests that the early interactions between spruces and fungi could potentially influence their growth trajectory.

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.

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.

Trade-offs between forest carbon stocks and harvests in a steady state - A multi-criteria analysis.

This paper provides a perspective for comparing trade-offs between harvested wood flows and forest carbon stocks with different forest management regimes. A constant management regime applied to a forest area with an even age-class distribution leads to a steady state, in which the annual harvest and carbon stocks remain constant over time. As both are desirable - carbon stocks for mitigating climate change and harvests for the economic use of wood and displacing fossil fuels - an ideal strategy should be chosen from a set of management regimes that are Pareto-optimal in the sense of multi-criteria decision-making. When choosing between Pareto-optimal alternatives, the trade-off between carbon stock and harvests is unavoidable. This trade-off can be described e.g. in terms of carbon payback times or carbon returns. As numerical examples, we present steady-state harvest levels and carbon stocks in a Finnish boreal forest region for different rotation periods, thinning intensities and collection patterns for harvest residues. In the set of simulated management practices, harvest residue collection presents the most favorable trade-off with payback times around 30-40 years; while Pareto-optimal changes in rotation or thinnings exhibited payback times over 100 years, or alternatively carbon returns below 1%. By extending the rotation period and using less-intensive thinnings compared to current practices, the steady-state carbon stocks could be increased by half while maintaining current harvest levels. Additional cases with longer rotation periods should be also considered, but were here excluded due to the lack of reliable data on older forest stands.

Early root growth and architecture of fast- and slow-growing Norway spruce (Picea abies) families differ-potential for functional adaptation.

The relationship between the growth rate of aboveground parts of trees and fine root development is largely unknown. We investigated the early root development of fast- and slow-growing Norway spruce (Picea abies (L.) H. Karst.) families at a developmental stage when the difference in size is not yet observed. Seedling root architecture data, describing root branching, were collected with the WinRHIZO™ image analysis system, and mixed models were used to determine possible differences between the two growth phenotypes. A new approach was used to investigate the spatial extent of root properties along the whole sample root from the base of 1-year-old seedlings to the most distal part of a root. The root architecture of seedlings representing fast-growing phenotypes showed ~30% higher numbers of root branches and tips, which resulted in larger root extensions and potentially a better ability to acquire nutrients. Seedlings of fast-growing phenotypes oriented and allocated root tips and biomass further away from the base of the seedling than those growing slowly, a possible advantage in nutrient-limited and heterogeneous boreal forest soils. We conclude that a higher long-term growth rate of the aboveground parts in Norway spruce may relate to greater allocation of resources to explorative roots that confers a competitive edge during early growth phases in forest ecosystems.

Functional-structural plant models: a growing paradigm for plant studies.

A number of research groups in various areas of plant biology as well as computer science and applied mathematics have addressed modelling the spatiotemporal dynamics of growth and development of plants. This has resulted in development of functional-structural plant models (FSPMs). In FSPMs, the plant structure is always explicitly represented in terms of a network of elementary units. In this respect, FSPMs are different from more abstract models in which a simplified representation of the plant structure is frequently used (e.g. spatial density of leaves, total biomass, etc.). This key feature makes it possible to build modular models and creates avenues for efficient exchange of model components and experimental data. They are being used to deal with the complex 3-D structure of plants and to simulate growth and development occurring at spatial scales from cells to forest areas, and temporal scales from seconds to decades and many plant generations. The plant types studied also cover a broad spectrum, from algae to trees. This special issue of Annals of Botany features selected papers on FSPM topics such as models of morphological development, models of physical and biological processes, integrated models predicting dynamics of plants and plant communities, modelling platforms, methods for acquiring the 3-D structures of plants using automated measurements, and practical applications for agronomic purposes.

Dynamics of leaf gas exchange, xylem and phloem transport, water potential and carbohydrate concentration in a realistic 3-D model tree crown.

BACKGROUND AND AIMS: Tree models simulate productivity using general gas exchange responses and structural relationships, but they rarely check whether leaf gas exchange and resulting water and assimilate transport and driving pressure gradients remain within acceptable physical boundaries. This study presents an implementation of the cohesion-tension theory of xylem transport and the Münch hypothesis of phloem transport in a realistic 3-D tree structure and assesses the gas exchange and transport dynamics. METHODS: A mechanistic model of xylem and phloem transport was used, together with a tested leaf assimilation and transpiration model in a realistic tree architecture to simulate leaf gas exchange and water and carbohydrate transport within an 8-year-old Scots pine tree. The model solved the dynamics of the amounts of water and sucrose solute in the xylem, cambium and phloem using a fine-grained mesh with a system of coupled ordinary differential equations. KEY RESULTS: The simulations predicted the observed patterns of pressure gradients and sugar concentration. Diurnal variation of environmental conditions influenced tree-level gradients in turgor pressure and sugar concentration, which are important drivers of carbon allocation. The results and between-shoot variation were sensitive to structural and functional parameters such as tree-level scaling of conduit size and phloem unloading. CONCLUSIONS: Linking whole-tree-level water and assimilate transport, gas exchange and sink activity opens a new avenue for plant studies, as features that are difficult to measure can be studied dynamically with the model. Tree-level responses to local and external conditions can be tested, thus making the approach described here a good test-bench for studies of whole-tree physiology.

Quantitative characterization of clumping in Scots pine crowns.

BACKGROUND AND AIMS: Proper characterization of the clumped structure of forests is needed for calculation of the absorbed radiation and photosynthetic production by a canopy. This study examined the dependency of crown-level clumping on tree size and growth conditions in Scots pine (Pinus sylvestris), and determined the ability of statistical canopy radiation models to quantify the degree of self-shading within crowns as a result of the clumping effect. METHODS: Twelve 3-D Scots pine trees were generated using an application of the LIGNUM model, and the crown-level clumping as quantified by the crown silhouette to total needle area ratio (STAR(crown)) was calculated. The results were compared with those produced by the stochastic approach of modelling tree crowns as geometric shapes filled with a random medium. KEY RESULTS: Crown clumping was independent of tree height, needle area and growth conditions. The results supported the capability of the stochastic approach in characterizing clumping in crowns given that the outer shell of the tree crown is well represented. CONCLUSIONS: Variation in the whole-stand clumping index is induced by differences in the spatial pattern of trees as a function of, for example, stand age rather than by changes in the degree of self-shading within individual crowns as they grow bigger.

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.

Predicting the decomposition of Scots pine, Norway spruce, and birch stems in Finland.

Models were developed for predicting the decomposition of dead wood for the main tree species in Finland, based on data collected from long-term thinning experiments in southern and central Finland. The decomposition rates were strongly related to the number of years after tree death. In contrast to previous studies, which have used the first-order exponential model, we found that the decomposition rate was not constant. Therefore, the Gompertz and Chapman-Richard's functions were fitted to the data. The slow initial decomposition period was mainly due to the fact that most dead trees remained standing as snags after their death. The initial period was followed by a period of rapid decomposition and, finally, by a period of moderately slow decomposition. Birch stems decomposed more rapidly than Scots pine and Norway spruce stems. Decomposition rates of Norway spruce stems were somewhat lower than those of Scots pine. Because the carbon concentration of decaying boles was relatively stable (about 50%) the rate of carbon loss follows that of mass loss. Models were also developed for the probability that a dead tree remains standing as a snag. During the first years after death, the probability was high. Thereafter, it decreased rapidly, the decrease being faster for birch stems than for Scots pine and Norway spruce stems. Almost all stems had fallen down within 40 years after their death. In Scots pine and Norway spruce, most snags remained hard and belonged to decay class 1. In birch, a higher proportion of snags belonged to the more advanced decay classes. The models provide a framework for predicting dead wood dynamics in managed as well as dense unthinned stands. The models can be incorporated into forest management planning systems, thereby facilitating estimates of carbon dynamics.

Analysis of biomass accumulation and stem size distributions over long periods in managed stands of Pinus sylvestris in Finland using the 3-PG model.

We tested the performance of a process-based model (PBM) in relation to long-term mensuration data from two sites in Finland where the stands were up to 90 years old and had been thinned at approximately 5-year intervals over the last 50 years. The PBM used was based on the 3-PG (Physiological Principles to Predict Growth) model developed by Landsberg and Waring (1997), with modifications in the biomass allocation routine, for which we used data and calculations by Vanninen (2003) to estimate the allocation coefficients and turnover rates. Site fertility was estimated in terms of known site-type characteristics. The model was evaluated in terms of stand development and its ability to simulate responses to thinning; stem numbers after thinning were specified at the dates when the thinning took place. Stand development in terms of basal area, volume and mean diameter at breast height, closely followed the measured characteristics of all stands. Foliage mass predictions were close to estimates obtained by an empirical method. The analysis shows that, under normal thinning regimes, a range of different thinning intensities can be adequately described using a simple multiplicative model relating the proportion of volume and foliage mass removed to the corresponding proportion of stem numbers. This model, together with stem allometry data, described the "growth" in mean diameter after thinning, which simply reflected the removal of the smaller trees. These results indicate that, with a single set of parameter values, 3-PG can provide good descriptions of the growth patterns of trees-in this case Pinus sylvestris L.-over long periods, including growth after repeated thinning. One of the outputs from the 3-PG model is mean stem diameter (B): we show that it is feasible to estimate stem size distributions, which changed considerably over the life of these stands, from B using the Weibull function. This shows that, given information about the Weibull parameters for particular species and cultural systems, it should be possible to use stem numbers and the B obtained from the 3-PG model to produce information about stem size distributions from simulated data.

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.

Application of volume growth and survival graphs in the evaluation of four process-based forest growth models.

Volume growth and survival (VGS) graphs, which show volume growth rate and risk of mortality for individual trees (or tree size classes), have been proposed as a tool for assessing the validity of models that describe the development over time of tree size distributions within forest stands. We examined the utility of the VGS method in evaluating four process-based models. The performance of the models FORSKA, 4C, MORG, and PipeQual is analyzed against long-term data from a Scots pine stand in Evo, Finland, and the models FORSKA and 4C are also assessed with respect to data from a beech stand in Fabrikschleichach, Germany. Comparison of the measurement-based VGS graphs with those produced from the model-based data shows that although the models yield similar stand-level predictions, they can differ widely in their projections of individual tree growth and size distributions. Examination of the discrepancies between models and data in the context of the VGS graphs reveals several areas in which the models could be improved. We conclude that the method is useful in model evaluation, especially if used in combination with indicators of stand structure, such as the height/diameter ratio.

Volume growth and survival graphs: a method for evaluating process-based forest growth models.

We investigated the relationships within forest stands between tree size and (a) stem volume growth rate and (b) risk of mortality for individual trees. Values of both x and y variables were plotted relative to the largest value in the stand. We refer to the resultant presentations as relative volume growth and relative survival graphs (VGSs). A pair of VGSs can be produced readily from an individual-tree growth model. It can also be constructed from consecutive sets of field measurements. Comparison of VGSs derived from model and measurement data provides a test of the validity of the components of the growth model. We have analyzed VGSs based on measurement data for Scots pine (Pinus sylvestris L.) in central Finland and for beech (Fagus sylvatica L.) in southern Germany. The graphs based on measurement data varied as a consequence of differences in competition, stand management, and tree species. We analyzed the relationship between VGSs and stand dynamics using a simple growth model. We found that different features of the VGSs imply characteristic tree size distributions in subsequent years. Thus, we conclude that if the VGSs generated by a model do not correspond to those based on field measurements, the model cannot be relied on to reproduce the development of tree-size distribution correctly. Relative growth and survival graphs thus provide a tool for evaluating complicated growth models.