A single-substrate model to interpret intra-annual stable isotope signals in tree-ring cellulose.

The carbon and oxygen stable isotope composition of wood cellulose (delta(13)C(cellulose) and delta(18)O(cellulose), respectively) reveal well-defined seasonal variations that contain valuable records of past climate, leaf gas exchange and carbon allocation dynamics within the trees. Here, we present a single-substrate model for wood growth to interpret seasonal isotopic signals collected in an even-aged maritime pine plantation growing in South-west France, where climate, soil and flux variables were also monitored. Observed seasonal patterns in delta(13)C(cellulose) and delta(18)O(cellulose) were different between years and individuals, and mostly captured by the model, suggesting that the single-substrate hypothesis is a good approximation for tree ring studies on Pinus pinaster, at least for the environmental conditions covered by this study. A sensitivity analysis revealed that the model was mostly affected by five isotopic discrimination factors and two leaf gas-exchange parameters. Modelled early wood signals were also very sensitive to the date when cell wall thickening begins (t(wt)). Our model could therefore be used to reconstruct t(wt) time series and improve our understanding of how climate influences this key parameter of xylogenesis.

Sink feedback regulation of photosynthesis in vines: measurements and a model.

An experimental and modelling study of source-sink interactions in Vitis vinifera L., cv. Cabernet Sauvignon, rooted cuttings under non-limiting environmental conditions with a 12 h photoperiod is presented here. After 4 h, measured photosynthesis, stomatal conductance and leaf carbohydrate content reached maximum values. Over the remainder of the photoperiod, photosynthesis and stomatal conductance decreased continuously, whereas leaf carbohydrate content remained relatively constant. Because the experiment took place in a non-limiting environment, the results suggest that stomatal regulation of photosynthesis was mediated by an internal factor, possibly related to sink activity. A simple 1-source, 2-sink model was developed to examine the extent to which the data could be explained by a hypothetical sink-to-source feedback mechanism mediated by carbohydrate levels in either the mesophyll, the source phloem or the phloem of one of the two sinks. Model simulations reproduced the data well under the hypothesis of a phloem-based feedback signal, although the data were insufficient to elucidate the detailed nature of such a signal. In a sensitivity analysis, the steady-state response of photosynthesis to sink activity was explored and predictions made for the partitioning of photosynthate between the two sinks. The analysis highlights the effectiveness of a phloem-based feedback signal in regulating the balance between source and sink activities. However, other mechanisms for the observed decline in photosynthesis, such as photoinhibition, endogenous circadian rhythms or hydraulic signals in the leaf cannot be excluded. Nevertheless, it is concluded that the phloem-based feedback model developed here may provide a useful working hypothesis for incorporation into plant growth models and for further development and testing.

Increased understanding of nutrient immobilization in soil organic matter is critical for predicting the carbon sink strength of forest ecosystems over the next 100 years.

The terrestrial biosphere is currently thought to be a significant sink for atmospheric carbon (C). However, the future course of this sink under rising [CO2] and temperature is uncertain. Some contrasting possibilities that have been suggested are: that the sink is currently increasing through CO2 fertilization of plant growth but will decline over the next few decades because of CO2 saturation and soil nutrient constraints; that the sink will continue to increase over the next century because rising temperature will stimulate the release of plant-available soil nitrogen (N) through increased soil decomposition; that, alternatively, the sink will not be sustained because the additional soil N released will be immobilized in the soil rather than taken up by plants; or that the sink will soon become negative because loss of soil C through temperature stimulation of soil respiration will override any CO2 or temperature stimulation of plant growth. Soil N immobilization is thus a key process; however, it remains poorly understood. In this paper we use a forest ecosystem model of plant-soil C and N dynamics to gauge the importance of this uncertainty for predictions of the future C sink of forests under rising [CO2] and temperature. We characterize soil N immobilization by the degree of variability of soil N:C ratios assumed in the model. We show that the modeled C sink of a stand of Norway spruce (Picea abies (L.) Karst.) in northern Sweden is highly sensitive to this assumption. Under increasing temperature, the model predicts a strong C sink when soil N:C is inflexible, but a greatly reduced C sink when soil N:C is allowed to vary. In complete contrast, increasing atmospheric [CO2] leads to a much stronger C sink when soil N:C is variable. When both temperature and [CO2] increase, the C sink strength is relatively insensitive to variability in soil N:C; significantly, however, with inflexible soil N:C the C sink is primarily a temperature response whereas with variable soil N:C, it is a combined temperature-CO2 response. Simulations with gradual increases of temperature and [CO2] indicate a sustained C sink over the next 100 years, in contrast to recent claims that the C sink will decline over the next few decades. Nevertheless, in using a relatively simple model, our primary aim is not to make precise predictions of the C sink over the next 100 years, but rather to highlight key areas of model uncertainty requiring further experimental clarification. Here we show that improved understanding of the processes underlying soil N immobilization is essential if we are to predict the future course of the forest carbon sink.

A simple model of light and water use evaluated for Pinus radiata.

An existing model of light and water use by crops (RESCAP) was adapted and evaluated for trees. In the model, growth on any given day is determined either by the amount of intercepted radiation (by means of the light utilization coefficient, epsilon) or by the maximum rate of water extraction by roots (a function of root biomass and soil water content). In either case, transpiration and growth are related by the water-use efficiency (q), which is inversely proportional to the daily mean saturation vapor pressure deficit (D). The model was applied to two Pinus radiata (D. Don) stands (control (C) and fertilized (F)) growing near Canberra, Australia, using data collected during the Biology of Forest Growth experiment (1983-1988). For both stands, predicted and measured soil water contents were in close agreement (r(2) > 0.9) over a 4-year period involving several wet-dry cycles. The parameter combination epsilon/qD was estimated to be 0.28 and 0.26 kg H(2)O (MJ total)(-1) kPa(-1) for the C and F stands, respectively. Because of the close physiological link between water use and CO(2) uptake, the results suggest that tree growth may be realistically simulated by simple models based on conservative values for epsilon and qD.

Carbon sequestration in the trees, products and soils of forest plantations: an analysis using UK examples.

A carbon-flow model for managed forest plantations was used to estimate carbon storage in UK plantations differing in Yield Class (growth rate), thinning regime and species characteristics. Time-averaged, total carbon storage (at equilibrium) was generally in the range 40-80 Mg C ha(-1) in trees, 15-25 Mg C ha(-1) in above- and belowground litter, 70-90 Mg C ha(-1) in soil organic matter and 20-40 Mg C ha(-1) in wood products (assuming product lifetime equalled rotation length). The rate of carbon storage during the first rotation in most plantations was in the range 2-5 Mg C ha(-1) year(-1).A sensitivity analysis revealed the following processes to be both uncertain and critical: the fraction of total woody biomass in branches and roots; litter and soil organic matter decomposition rates; and rates of fine root turnover. Other variables, including the time to canopy closure and the possibility of accelerated decomposition after harvest, were less critical. The lifetime of wood products was not critical to total carbon storage because wood products formed only a modest fraction of the total.The average increase in total carbon storage in the tree-soil-product system per unit increase in Yield Class (m(3) ha(-1) year(-1)) for unthinned Picea sitchensis (Bong.) Carr. plantations was 5.6 Mg C ha(-1). Increasing the Yield Class from 6 to 24 m(3) ha(-1) year(-1) increased the rate of carbon storage in the first rotation from 2.5 to 5.6 Mg C ha(-1) year(-1) in unthinned plantations. Thinning reduced total carbon storage in P. sitchensis plantations by about 15%, and is likely to reduce carbon storage in all plantation types.If the objective is to store carbon rapidly in the short term and achieve high carbon storage in the long term, Populus plantations growing on fertile land (2.7 m spacing, 26-year rotations, Yield Class 12) were the best option examined. If the objective is to achieve high carbon storage in the medium term (50 years) without regard to the initial rate of storage, then plantations of conifers of any species with above-average Yield Classes would suffice. In the long term (100 years), broadleaved plantations of oak and beech store as much carbon as conifer plantations. Mini-rotations (10 years) do not achieve a high carbon storage.

Predicted changes in the synchrony of larval emergence and budburst under climatic warming.

The impact of climatic warming on the synchrony of insect and plant phenologies was modelled in the case of winter moth (Operophtera brumata) and Sitka spruce (Picea sitchensis) in the Scottish uplands. The emergence of winter moth larvae was predicted with a thermal time requirement model and the budburst of Sitka spruce was predicted from a previously published model (Cannell and Smith 1983) based on winter chilling and thermal time. The date of emergence of winter moth larvae was predicted to occur earlier under climatic warming but the date of budburst of Sitka spruce was not greatly changed, resulting in decreased synchrony between larval emergence and budburst. The general question of how a change of climate might affect phenological synchrony and insect abundance is discussed.

Analytical model of carbon storage in the trees, soils, and wood products of managed forests.

The carbon balance between managed forests and the atmosphere depends critically on the frequency and intensity of harvesting, and the lifetime of harvested products. To assess more quantitatively the nature of this dependence, a theoretical analysis, previously applied to carbon storage in trees and wood products only, is extended here to include the carbon in forest floor detritus and soil. A dimensionless combination of the parameters of the model, alpha, with critical value alpha(c), is identified such that for alpha < alpha(c), the conversion of old-growth forest to managed forest releases carbon to the atmosphere in the long term. Parameter alpha is given by the combination f(t)D/T(*), where f(t) is the fraction of old-growth forest carbon stored in trees, D is the residence time of harvested biomass (wood products and slash debris) within the system, and T(*) is the rotation period for maximum sustained yield (maximum mean annual increment). The critical value alpha(c), typically in the range 0.5-0.7, is derived for a variety of forest types. Parameter alpha determines the degree to which the carbon accumulated in harvested biomass offsets the loss of carbon in trees due to felling and in soils due to reduced litter input. When alpha > alpha(c), long-term carbon storage is optimized by harvesting for maximum sustained yield.

A model of carbon storage in forests and forest products.

This paper discusses the general formulation of a model that describes carbon storage in a forest and its timber products as a function of the forest growth curve, the rotation period and the carbon retention curves for the timber products. After a number of rotations, the rotation-averaged quantity of stored carbon approaches an asymptotic value. It is shown that, when forests are managed for maximum sustained yield of biomass, the contribution to asymptotic carbon storage from timber products is about 2.5D/T* times the contribution from living trees, where D is the characteristic decay time for reconversion of timber products to carbon dioxide, and T* is the normal rotation period for maximum sustained yield. For a given value of D/T*, carbon storage can be optimized if the policy of maximizing sustained yield is relaxed. For D/T* < 1, as the rotation period is increased indefinitely, the asymptotic level of carbon storage increases monotonically toward the value of the carbon content of living trees at maturity, g(f). For D/T* > 1, there is a finite, optimal rotation period, T(o), greater than T*, for which asymptotic carbon storage is greater than g(f). As D/T* tends to large values, however, T(o) tends to T*, so that, in this limit, management for maximum sustained yield also ensures maximum carbon storage. From initial planting, the time taken to reach asymptotic carbon storage decreases as the normal rotation period, T*, decreases, but increases almost linearly with increasing decay time of timber products, D. This result qualifies the short-term value of any particular planting strategy.