Water relations in tree physiology: where to from here?

We look back over 50 years of research into the water relations of trees, with the objective of assessing the maturity of the topic in terms of the idea of a paradigm, put forward by Kuhn in 1962. Our brief review indicates that the physical processes underlying the calculation of transpiration are well understood and accepted, and knowledge of those processes can be applied if information about the leaf area of trees, and stomatal conductance, is available. Considerable progress has been made in understanding the factors governing stomatal responses to environment, with insights into how the hydraulic conducting system of trees determines the maximum aperture of stomata. Knowledge about the maximum stomatal conductance values likely to be reached by different species, and recognition that stomatal responses to increasing atmospheric vapor pressure deficits are in fact responses to water loss from leaves, provides the basis for linking these responses to information about hydraulic conductance through soil­root­stem­branch systems. Improved understanding in these areas is being incorporated into modern models of stomatal conductance and responses to environmental conditions. There have been significant advances in understanding hydraulic pathways, including cavitation and its implications. A few studies suggest that the major resistances to water flux within trees are not in the stem but in the branches. This insight may have implications for productivity: it may be advantageous to select trees with the genetic propensity to produce short branches in stands with open canopies. Studies on the storage of water in stems have provided improved understanding of fluxes from sapwood at different levels. Water stored in the stems of large trees may provide up to 20­30% daily sap flow, but this water is likely to be replaced by inflows at night. In dry conditions transpiration by large trees may be maintained from stored water for up to a week, but flows from storage may be more important in refilling cavitated xylem elements and hence ensuring that the overall hydraulic conductivity of stems is not reduced. Hydraulic redistribution of water in the soil may make a contribution to facilitating root growth in dry soils and modifying resource availability. We conclude that the field of tree water relations is mature, in the sense that the concepts underlying models describing processes and system responses to change are well-tested and accepted and there are few, if any, serious anomalies emerging. Models are essentially formal statements about the way we think systems work. They are always subject to further testing, refinement and improvements. Gaps in knowledge appear within the framework of accepted concepts and mechanisms research is needed to fill those gaps. The models currently available can be used to scale estimates of transpiration from leaf to landscape levels and predict species responses to drought. The focus in tree water relations has shifted to examine the climatic thresholds at which drought, high temperatures and vapor pressure deficits cause mortality. Tree death may be caused by hydraulic collapse following irreversible cavitation or extremely low water potentials, but recent research indicates that the relative sensitivity of stomatal conductance and whole-plant hydraulic conductance plays a major role in determining plant responses to drought.

Does night-time transpiration contribute to anisohydric behaviour in a Vitis vinifera cultivar?

The hypothesis that vines of the Semillon wine grape variety show anisohydric behaviour was tested, i.e. that tissue hydration is unstable under fluctuating environmental conditions. Stomatal conductance and transpiration rates from leaves were measured during the day and at night. Leaf water potential (Psi(l)) in Semillon was negatively correlated to vapour pressure deficit (VPD) both predawn and during the day. Furthermore, Psi(l) fell to significantly lower values than in any of the nine other varieties examined. Night-time values of stomatal conductance (g(n)) and transpiration (E(n)) in Semillon were up to four times higher than in other varieties; plants enclosed in plastic bags overnight to reduce E(n) resulted in better plant-soil equilibration so that predawn Psi(l) in Semillon was the same as in Grenache. These data indicate that the hypothesis is supported, and that night-time transpiration contributes significantly to the low Psi(l) values in Semillon during warm, dry nights. The other contributing factor is daytime stomatal conductance (g(day)), which in Semillon leaves was higher than in other varieties, although the decline in g(day) with increasing VPD was greater in Semillon than in Shiraz or Grenache. The high values of g(day) were associated with high rates of transpiration (E(day)) by Semillon through a day when VPD reached 4.5 kPa. When compared to other varieties, Semillon was not unusual in terms of root length density, stomatal density, xylem sap abscisic acid, or leaf electrolyte leakage. Night-time and daytime water loss and insufficient stomatal regulation therefore account for the tendency to anisohydric behaviour shown by Semillon.

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.

Conversion of canopy intercepted radiation to photosynthate: review of modelling approaches for regional scales.

A fundamental component of most models of terrestrial carbon balance is an estimate of plant canopy photosynthetic uptake driven by radiation interception by the canopy. In this article, we review approaches used to model the conversion of radiation into photosynthate. As this process is well understood at the leaf-scale, the modelling problem is essentially one of up-scaling, to canopy, regional or global scale. Our review therefore focuses on issues of scaling, including model identification, parameterisation and validation at large scales. Four different approaches are commonly taken to modelling photosynthate production at large scales: the maximum productivity, resource-use efficiency, big-leaf, and sun-shade models. Models representing each of these approaches are discussed and model predictions compared with estimates of gross primary productivity derived from eddy covariance data measured above a Sitka spruce forest. The sun-shade model was found to perform best at all time scales considered. However, other models had significant advantages including simplicity of implementation and the ability to combine the model with remotely-sensed information on vegetation radiation interception. We conclude that all four approaches can be successfully used to model photosynthetic uptake and that the best approach in a given situation will depend on model objectives and data availability.

Corrigendum to: Conversion of canopy intercepted radiation to photosynthate: a review of modelling approaches for regional scales.

A fundamental component of most models of terrestrial carbon balance is an estimate of plant canopy photosynthetic uptake driven by radiation interception by the canopy. In this article, we review approaches used to model the conversion of radiation into photosynthate. As this process is well understood at the leaf-scale, the modelling problem is essentially one of up-scaling, to canopy, regional or global scale. Our review therefore focuses on issues of scaling, including model identification, parameterisation and validation at large scales. Four different approaches are commonly taken to modelling photosynthate production at large scales: the maximum productivity, resource-use efficiency, big-leaf, and sun-shade models. Models representing each of these approaches are discussed and model predictions compared with estimates of gross primary productivity derived from eddy covariance data measured above a Sitka spruce forest. The sun-shade model was found to perform best at all time scales considered. However, other models had significant advantages including simplicity of implementation and the ability to combine the model with remotely-sensed information on vegetation radiation interception. We conclude that all four approaches can be successfully used to model photosynthetic uptake and that the best approach in a given situation will depend on model objectives and data availability.

Analysis of the growth of rimu (Dacrydium cupressinum) in South Westland, New Zealand, using process-based simulation models.

Two process-based models were used to identify the environmental variables limiting productivity in a pristine, mature forest dominated by rimu (Dacrydium cupressinum Sol. ex Lamb.) trees in South Westland, New Zealand. A model of canopy net carbon uptake, incorporating routines for radiation interception, photosynthesis and water balance was used to determine a value for quantum efficiency when climate variables were not limiting. The annual net carbon uptake by the canopy was estimated to be 1.1 kg C m(-2) and the quantum efficiency 22.6 mmol mol quanta(-1). This value of quantum efficiency, combined with other parameters obtainable from the literature, was then used in a model of forest productivity (3-PG), to simulate changes in net productivity and the allocation of carbon to tree components. The model was adjusted to match a measured stem increment of 10.6 Mg ha(-1) over a period of 13 years. To achieve this while maintaining a low, but stable value for leaf area index, it was necessary to set the site fertility rating very low and select high values for the parameters describing the proportional allocation of total carbon to roots. This approach highlighted nutrient availability as the principal constraint on productivity for the ecosystem and identified critical measurements that will be necessary for using the model to predict the effects of climate change on carbon sequestration. The low rates of carbon uptake and productivity are consistent with the low nutrient supply available from the highly leached, acid soils, most likely attributable to frequent saturation and a very shallow aerobic zone.

Process-based models for forest ecosystem management: current state of the art and challenges for practical implementation.

Recent progress toward the application of process-based models in forestmanagement includes the development of evaluation and parameter estimation methods suitable for models with causal structure, and the accumulation of data that can be used in model evaluation. The current state of the art of process modeling is discussed in the context of forest ecosystem management. We argue that the carbon balance approach is readily applicable for projecting forest yield and productivity, and review several carbon balance models for estimating stand productivity and individual tree growth and competition. We propose that to develop operational models, it is necessary to accept that all models may have both empirical and causal components at the system level. We present examples of hybrid carbon balance models and consider issues that currently require incorporation of empirical information at the system level. We review model calibration and validation methods that take account of the hybrid character of models. The operational implementation of process-based models to practical forest management is discussed. Methods of decision-making in forest management are gradually moving toward a more general, analytical approach, and it seems likely that models that include some process-oriented components will soon be used in forestry enterprises. This development is likely to run parallel with the further development of ecophysiologically based models.