Modelling the response of tree growth to temperature and CO2 elevation as related to the fertility and current temperature sum of a site

A methodology for simulating climate change impacts on tree growth was introduced into a statistical growth and yield model in relation to variations in site fertility and location implemented with current temperature sum. This was based on a procedure in which the relative enhancement in stem volume growth was calculated from short-term runs of a physiological simulation model for Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies (L.) Karst.) and silver birch (Betula pendula Roth.) stands. These simulations were made for a set of stands with species-specific variations in stand characteristics, location and fertility type first in current climatic conditions and then in different combinations of CO₂ and temperature elevations. Based on these simulations, the relative enhancement of volume growth induced by the climate change (relative scenario effect, RSEv) was calculated and modelled in relation to: (i) CO₂ and temperature elevation, stand density and the competition status of the tree in its stand, and (ii) variations in site fertility type and current temperature sum of a stand. Finally, these transfer functions for RSEv were applied to adapt the stem volume growth in the statistical growth and yield model to reflect the response to climate change.     

Effects of seawater pCO2 and temperature on shell growth, shell stability, condition and cellular stress of Western Baltic Sea Mytilus edulis (L.) and Arctica islandica (L.)

Acidification of the World’s oceans may directly impact reproduction, performance and shell formation of marine calcifying organisms. In addition, since shell production is costly and stress in general draws on an organism’s energy budget, shell growth and stability of bivalves should indirectly be affected by environmental stress. The aim of this study was to investigate whether a combination of warming and acidification leads to increased physiological stress (lipofuscin accumulation and mortality) and affects the performance [shell growth, shell breaking force, condition index (C_i)] of young Mytilus edulis and Arctica islandica from the Baltic Sea. We cultured the bivalves in a fully-crossed 2-factorial experimental setup (seawater (sw) pCO₂ levels “low”, “medium” and “high” for both species, temperature levels 7.5, 10, 16, 20 and 25 °C for M. edulis and 7.5, 10 and 16 °C for A. islandica) for 13 weeks in summer. Mytilus edulis and A. islandica appeared to tolerate wide ranges of sw temperature and pCO₂. Lipofuscin accumulation of M. edulis increased with temperature while the C_i decreased, but shell growth of the mussels only sharply decreased while its mortality increased between 20 and 25 °C. In A. islandica, lipofuscin accumulation increased with temperature, whereas the C_i, shell growth and shell breaking force decreased. The pCO₂ treatment had only marginal effects on the measured parameters of both bivalve species. Shell growth of both bivalve species was not impaired by under-saturation of the sea water with respect to aragonite and calcite. Furthermore, independently of water temperatures shell breaking force of both species and shell growth of A. islandica remained unaffected by the applied elevated sw pCO₂ for several months. Only at the highest temperature (25 °C), growth arrest of M. edulis was recorded at the high sw pCO₂ treatment and the C_i of M. edulis was slightly higher at the medium sw pCO₂ treatment than at the low and high sw pCO₂ treatments. The only effect of elevated sw pCO₂ on A. islandica was an increase in lipofuscin accumulation at the high sw pCO₂ treatment compared to the medium sw pCO₂ treatment. Our results show that, despite this robustness, growth of both M. edulis and A. islandica can be reduced if sw temperatures remain high for several weeks in summer. As large body size constitutes an escape from crab and sea star predation, this can make bivalves presumably more vulnerable to predation—with possible negative consequences on population growth. In M. edulis, but not in A. islandica, this effect is amplified by elevated sw pCO₂. We follow that combined effects of elevated sw pCO₂ and ocean warming might cause shifts in future Western Baltic Sea community structures and ecosystem services; however, only if predators or other interacting species do not suffer as strong from these stressors.     

More asymmetric tree competition brings about more evapotranspiration and less runoff from the forest ecosystems: A simulation study

The present paper reports how stand size-structure dynamics due to competition between different-sized trees affect long-term forested water balance in Japanese cool-temperate planted stands (evergreen coniferous Cryptomeria japonica and deciduous coniferous Larix kaempferi stands) using a fully coupled multi-layered meteorological surface physics—terrestrial ecosystems model. The simulation captured the well-known annual variation in leaf area index (LAI) accurately with stand age in monocultured and even-aged stands; the occurrence of maximum LAI during the early growth stage and then a gradual decline followed by a steady state after the maximum LAI. The simulations also detected a high dependency of annual evapotranspiration (AETr) on LAI with stand age that is well known by prior observational researches. In the C. japonica (shade-tolerant late-successional species) stand, the relationship between annual net primary productivity of an individual tree (NPP_ind) and individual tree mass (w) changed from linear to a convex curve during self-thinning, indicating that the degree of asymmetric tree competition intensified with forest stand development. The higher degree of competitive asymmetry characterized by the convex-shaped NPPind-w relationship produced greater size inequality, i.e., the formation of trees stratified by height. Under such conditions, AETr and annual transpiration (ATr) were mainly regulated by larger trees. On the other hand, the NPPind-w relationships in the L. kaempferi (shade-intolerant early-successional species) stand were linear throughout the simulated period, indicating the lower degree of competitive asymmetry. Under such conditions, the growth of intermediate-sized trees was enhanced and these trees became a dominant source of AETr (and also ATr) during self-thinning. Furthermore, the sensitivity analysis of the effects of ecophysiological parameters such as foliage profile (i.e., vertical distribution of leaf area density) of an individual tree (distribution pattern is described by the parameter η), the maximum carboxylation velocity (V_cmax0) and biomass allocation pattern of individual plant growth (μ₁) on AETr, ATr and annual runoff (AR_off) showed that the temporal trends of AETr, ATr, AR_off and NPPind-w relationships were completely the same as those in the control simulations. However, the NPPind-w relationship during self-thinning indicated higher degrees of competitive asymmetry when η or V_cmax0 were greater than those in the control simulation and generated greater AETr and ATr and thus smaller AR_off. We found that more asymmetric tree competition brings about greater size inequality between different-sized trees and thus more evapotranspiration and less runoff in a forest stand. Overall, our simulation approach revealed that not only LAI dynamics but also plant competition, and thus size-structure dynamics, in a forest ecosystem are essential to long-term future projections of forested water balance.     

ANAFORE: A stand-scale process-based forest model that includes wood tissue development and labile carbon storage in trees

A stand-scale forest model has been developed that dynamically simulates, besides carbon (C) and water (H₂O) fluxes, wood tissue development from physiological principles. The forest stand is described as consisting of trees of different size cohorts (for example, dominant, co-dominant and suppressed trees), either of the same or of different species (deciduous or coniferous). Half-hourly C and H₂O fluxes are modeled at the leaf, tree and stand level. In addition to total growth and yield, the model simulates the daily evolution of tracheid or vessel biomass and radius, parenchyma and branch development. From these data early and latewood biomass, wood tissue composition and density are calculated. Simulation of the labile C stored in the living tissues allows for simulation of trans-seasonal and trans-yearly effects, and improved simulations of long-term effects of environmental stresses on growth. A sensitivity analysis was performed to indicate the main parameters influencing simulated stem growth and wood quality at the tree and stand level. Case studies were performed for a temperate pine forest to illustrate the main model functioning and, more in particular, the simulation of the wood quality. The results indicate that the ANAFORE model is a useful tool for simultaneous analyses of wood quality development and forest ecosystem functioning.     

Mathematical analysis of change in forest carbon use efficiency with stand development: A case study on Abies veitchii Lindl.

Changes in carbon use efficiency (CUE), which is defined as the ratio of net primary production (NPP) to gross primary production (GPP), were analyzed for Abies veitchii Lindl. forests with respect to stand development by developing a simple mathematical model incorporating data on physiological variables and leaf mass ratio. A decrease in CUE with stand development was successfully expressed as a function of stand biomass (y) based on the following three assumptions: (1) a power-law relationship between mean respiration and mean individual tree mass, (2) a power-functional relationship between mean gross primary production and mean individual tree mass, and (3) self-thinning relationship between stand biomass and density. Based on this model, a parameter of CUE–y relationship was defined, and it was clarified that CUE decrease with stand development is caused not by the ratio of specific respiration rate to specific gross photosynthetic rate, but by leaf mass ratio. Since CUE is high in young forests, helpful information on selecting woody species when planting seedlings was provided from the viewpoints of reducing CO₂ in the atmosphere and global warming.     

Response of tree growth to a changing climate in boreal central Canada: A comparison of empirical,process-based,and hybrid modelling approaches

The impact of 2 × CO₂ driven climate change on radial growth of boreal tree species Pinus banksiana Lamb., Populus tremuloides Michx. and Picea mariana (Mill.) BSP growing in the Duck Mountain Provincial Forest of Manitoba (DMPF), Canada, is simulated using empirical and process-based model approaches. First, empirical relationships between growth and climate are developed. Stepwise multiple-regression models are conducted between tree-ring growth increments (TRGI) and monthly drought, precipitation and temperature series. Predictive skills are tested using a calibration–verification scheme. The established relationships are then transferred to climates driven by 1× and 2 × CO₂ scenarios using outputs from the Canadian second-generation coupled global climate model. Second, empirical results are contrasted with process-based projections of net primary productivity allocated to stem development (NPP_s). At the finest scale, a leaf-level model of photosynthesis is used to simulate canopy properties per species and their interaction with the variability in radiation, temperature and vapour pressure deficit. Then, a top-down plot-level model of forest productivity is used to simulate landscape-level productivity by capturing the between-stand variability in forest cover. Results show that the predicted TRGI from the empirical models account for up to 56.3% of the variance in the observed TRGI over the period 1912–1999. Under a 2 × CO₂ scenario, the predicted impact of climate change is a radial growth decline for all three species under study. However, projections obtained from the process-based model suggest that an increasing growing season length in a changing climate could counteract and potentially overwhelm the negative influence of increased drought stress. The divergence between TRGI and NPP_s simulations likely resulted, among others, from assumptions about soil water holding capacity and from calibration of variables affecting gross primary productivity. An attempt was therefore made to bridge the gap between the two modelling approaches by using physiological variables as TRGI predictors. Results obtained in this manner are similar to those obtained using climate variables, and suggest that the positive effect of increasing growing season length would be counteracted by increasing summer temperatures. Notwithstanding uncertainties in these simulations (CO₂ fertilization effect, feedback from disturbance regimes, phenology of species, and uncertainties in future CO₂ emissions), a decrease in forest productivity with climate change should be considered as a plausible scenario in sustainable forest management planning of the DMPF.     

The physiological and molecular responses of larvae from the reef-building coral Pocillopora damicornis exposed to near-future increases in temperature and pCO2

Given the threats of greenhouse gas emissions and a changing climate to marine ecosystems, there is an urgent need to better understand the response of not only adult corals, which are particularly sensitive to environmental changes, but also their larvae, whose mechanisms of acclimation to both temperature increases and ocean acidification are not well understood. Brooded larvae from the reef coral Pocillopora damicornis collected from Nanwan Bay, Southern Taiwan, were exposed to ambient or elevated temperature (25 or 29 °C) and pCO₂ (415 or 635 μatm) in a factorial experiment for 9 days, and a variety of physiological and molecular parameters were measured. Respiration and rubisco protein expression decreased in larvae exposed to elevated temperature, while those incubated at high pCO₂ were larger in size. Collectively, these findings highlight the complex metabolic and molecular responses of this life history stage and the need to integrate our understanding across multiple levels of biological organization. Our results also suggest that for this pocilloporid larval life stage, the impacts of elevated temperature are likely a greater threat under near-future predictions for climate change than ocean acidification.     

Application of a three-dimensional model for assessing effects of small clear-cuttings on radiation and soil temperature

A three-dimensional model Mixfor-3D of soil–vegetation–atmosphere transfer (SVAT) was developed and applied to estimate possible effects of tree clear-cutting on radiation and soil temperature regimes of a forest ecosystem. The Mixfor-3D model consists of several closely coupled 3D sub-models describing: forest stand structure; radiative transfer in a forest canopy; turbulent transfer of sensible heat, H₂O and CO₂ between ground surface and the atmospheric surface layer; evapotranspiration of ground surface vegetation and soil; heat and moisture transfer in soil. The model operates with the horizontal grid resolution, 2 m × 2 m; vertical resolution, 1 m and primary time step, 1 h.     

Tolerance of juvenile barnacles (Amphibalanus improvisus) to warming and elevated pCO2

We investigated the impacts of warming and elevated pCO₂ on newly settled Amphibalanus improvisus from Kiel Fjord, an estuarine ecosystem characterized by significant natural pCO₂ variability. In two experiments, juvenile barnacles were maintained at two temperature and three pCO₂ levels (20/24 °C, 700–2,140 μatm) for 8 weeks in a batch culture and at four pCO₂ levels (20 °C, 620–2,870 μatm) for 12 weeks in a water flow-through system. Warming as well as elevated pCO₂ hardly affected growth or the condition index of barnacles, although some factor combinations led to temporal significances in enhanced or reduced growth with an increase in pCO₂. While warming increased the shell strength of A. improvisus individuals, elevated pCO₂ had only weak effects. We demonstrate a strong tolerance of juvenile A. improvisus to mean acidification levels of about 1,000 μatm pCO₂ as is already naturally experienced by the investigated barnacle population.     

Ocean acidification and warming alter photosynthesis and calcification of the symbiont-bearing foraminifera Marginopora vertebralis

The impact of elevated CO₂ and temperature on photosynthesis and calcification in the symbiont-bearing benthic foraminifer Marginopora vertebralis was studied. Individual specimens of M. vertebralis were collected from Heron Island on the southern Great Barrier Reef (Australia). They were maintained for 5 weeks at different temperatures (28, 32 °C) and pCO₂ (400, 1,000 µatm) levels spanning a range of current and future climate-change scenarios. The photosynthetic capacity of M. vertebralis was measured with O₂ microsensors and a pulse-amplitude-modulated chlorophyll (Chl) fluorometer, in combination with estimates of Chl a and Chl c ₂ concentrations and calcification rates. After 5 weeks, control specimens remained unaltered for all parameters. Chlorophyll a concentrations significantly decreased in the specimens at 1,000 µatm CO₂ for both temperatures, while no change in Chl c ₂ concentration was observed. Photoinhibition was observed under elevated CO₂ and temperature, with a 70–80 % decrease in the maximum quantum yield of PSII. There was no net O₂ production at elevated temperatures in both CO₂ treatments as compared to the control temperature, supporting that temperature has more impact on photosynthesis and O₂ flux than changes in ambient CO₂. Photosynthetic pigment loss and a decrease in photochemical efficiency are thus likely to occur with increased temperature. The elevated CO₂ and high temperature treatment also lead to a reduction in calcification rate (from +0.1 to >?0.1 % day^?1). Thus, both calcification and photosynthesis of the major sediment-producing foraminifer M. vertebralis appears highly vulnerable to elevated temperature and ocean acidification scenarios predicted in climate-change models.     

Using a biogeochemistry model in simulating forests productivity responses to climatic change and [CO2] increase: example of Pinus halepensis in Provence (south-east France)

Tree-ring chronologies provide long-term records of growth in natural environmental conditions and may be used to evaluate the impacts of climatic change and [CO₂] increase on forest productivity. This study focuses on 21 Pinus halepensis forest stands in calcareous Provence (in the south-east France). Changes in productivity are simulated using the global biogeochemistry model BIOME3, that we have adapted to run with chronological data. Tree-ring data (width and density) were used to estimate, for each stand, an observed series of changes in productivity. Simulated and observed productivity changes are then compared to validate the chronological biogeochemistry model BIOME3C. Variations in productivity were well reconstructed at 15 sites. After this validation, BIOME3C was used to simulate forest productivity changes for a 2×CO₂ scenario. The 2×CO₂ climate used as input was obtained using results from Météo-France’s ARPEGE atmospheric general circulation model (AGCM), downscaled to local meteorological stations. Productivity increases moderately for all stands (from 17 to 24%) when climatic changes alone were taken into account. The main factor responsible for this increase is a reduction in summer drought severity. Productivity increases highly for all stands (from 72 to 86%) when the physiological fertilising effect of the [CO₂] increase is considered separately. When both climatic changes and the [CO₂] increase were taken into account, productivity increases highly, from 107% (for Moustier) to 141% (for La Ciotat). The direct fertilising effect of [CO₂] increase has a greater influence on the forest stands productivity than the indirect climatic changes effect. These results also exhibit the importance of the synergy between the effects of climate change and [CO₂] increase, as the increase in productivity resulting from the combined effects are more than the sum of the individual CO₂ and climate effects. Although the detected effects of global change during the 20th century were slight, acceleration of these changes is likely to lead to great changes in the future productivity of P. halepensis forests.     

Water and carbon fluxes above European coniferous forests modelled with artificial neural networks

Artificial neural networks are used to select a minimal set of input variables to model water vapour and carbon exchange of coniferous forest ecosystems, independently of tree species and without detailed physiological information. Neural networks are used because of their power to fit highly non-linear relations between input and output-variables. Radiation, temperature, vapour pressure deficit and time of the day showed to be the dynamic input variables that determine ecosystem water fluxes. The same variables, together with projected leaf area index are needed for modelling CO₂-fluxes. The results for the individual sites show that the neural networks found mean water and carbon flux responses to the driving variables valid for all sites. The sensitivity analysis of the derived neural networks shows that the LAI-effect of the CO₂-flux model is overfitted because of the low variability of LAI. However, the predictions of CO₂-fluxes of sites not included in the calibration set indicate that the LAI-response of the network is reliable and that results can be used as a first estimate of the net ecosystem carbon exchange of the forest sites. Independent predictions of forest ecosystem vapour fluxes were equally satisfying as empirical models specifically calibrated for the individual sites. The results indicate that both short term water and carbon fluxes of European coniferous forests can be modelled without using detailed physiological and site specific information.     

Foliage profiles of individual trees determine competition,self-thinning,biomass and NPP of a Cryptomeria japonica forest stand: A simulation study based on a stand-scale process-based forest model

A simulation study was carried out to investigate simultaneously the effects of eco-physiological parameters on competitive asymmetry, self-thinning, stand biomass and NPP in a temperate forest using an atmosphere–vegetation dynamics interactive model (MINoSGI). In this study, we selected three eco-physiological relevant parameters as foliage profiles (i.e. vertical distribution of leaf area density) of individual trees (distribution pattern is described by the parameter η), biomass allocation pattern in individual tree growth (χ) and the maximum carboxylation velocity (V_max). The position of the maximal leaf area density shifts upward in the canopy with increasing η. For scenarios with η < 4 (foliage concentrated in the lowest canopy layer) or η > 12 (foliage concentrated in the uppermost canopy layer), a low degree of competitive asymmetry was produced. These scenarios resulted in the survival of subordinate trees due to a brighter lower canopy environment when η < 4 or the generation of spatially separated foliage profiles between dominant and subordinate trees when η > 12. In contrast, competition between trees was most asymmetric when 4 ≤ η ≤ 12 (vertically widespread foliage profile in the canopy), especially when η = 8. In such cases, vertically widespread foliage of dominant trees lowered the opportunity of light acquisition for subordinate trees and reduced their carbon gain. The resulting reduction in carbon gain of subordinate trees yielded a higher degree of competitive asymmetry and ultimately higher mortality of subordinate trees. It was also shown that 4 ≤ η ≤ 12 generated higher self-thinning speed, smaller accumulated NPP, litter-fall and potential stand biomass as compared with the scenarios with η < 4 or η > 12. In contrast, our simulation revealed small effects of χ or V_max on the above-mentioned variables as compared with those of η. In particular, it is notable that greater V_max would not produce greater potential stand biomass and accumulated NPP although it has been thought that physiological parameters relevant to photosynthesis such as V_max influence dynamic changes in forest stand biomass and NPP (e.g. the greater the V_max, the greater the NPP). Overall, it is suggested that foliage profiles rather than biomass allocation or maximum carboxylation velocity greatly govern forest dynamics, stand biomass, NPP and litter-fall.     

Tolerance of Hyas araneus zoea I larvae to elevated seawater PCO2 despite elevated metabolic costs

Early life stages of marine crustaceans respond sensitively to elevated seawater PCO₂. However, the underlying physiological mechanisms have not been studied well. We therefore investigated the effects of elevated seawater PCO₂ on oxygen consumption, dry weight, elemental composition, median developmental time (MDT) and mortality in zoea I larvae of the spider crab Hyas araneus (Svalbard 79°N/11°E; collection, May 2009; hatch, December 2009). At the time of moulting, oxygen consumption rate had reached a steady state level under control conditions. In contrast, elevated seawater PCO₂ caused the metabolic rate to rise continuously leading to a maximum 1.5-fold increase beyond control level a few days before moulting into the second stage (zoea II), followed by a pronounced decrease. Dry weight of larvae reared under high CO₂ conditions was lower than in control larvae at the beginning of the moult cycle, yet this difference had disappeared at the time of moulting. MDT of zoea I varied between 45 ± 1 days under control conditions and 42 ± 2 days under the highest seawater CO₂ concentration. The present study indicates that larval development under elevated seawater PCO₂ levels results in higher metabolic costs during premoulting events in zoea I. However, H. araneus zoea I larvae seem to be able to compensate for higher metabolic costs as larval MDT and survival was not affected by elevated PCO₂ levels.     

Long-term and trans-life-cycle effects of exposure to ocean acidification in the green sea urchin Strongylocentrotus droebachiensis

Anthropogenic CO₂ emissions are acidifying the world’s oceans. A growing body of evidence demonstrates that ocean acidification can impact survival, growth, development and physiology of marine invertebrates. Here, we tested the impact of long-term (up to 16 months) and trans-life-cycle (adult, embryo/larvae and juvenile) exposure to elevated pCO₂ (1,200 μatm, compared to control 400 μatm) on the green sea urchin Strongylocentrotus droebachiensis. Female fecundity was decreased 4.5-fold when acclimated to elevated pCO₂ for 4 months during reproductive conditioning, while no difference was observed in females acclimated for 16 months. Moreover, adult pre-exposure for 4 months to elevated pCO₂ had a direct negative impact on subsequent larval settlement success. Five to nine times fewer offspring reached the juvenile stage in cultures using gametes collected from adults previously acclimated to high pCO₂ for 4 months. However, no difference in larval survival was observed when adults were pre-exposed for 16 months to elevated pCO₂. pCO₂ had no direct negative impact on juvenile survival except when both larvae and juveniles were raised in elevated pCO₂. These negative effects on settlement success and juvenile survival can be attributed to carry-over effects from adults to larvae and from larvae to juveniles. Our results support the contention that adult sea urchins can acclimate to moderately elevated pCO₂ in a matter of a few months and that carry-over effects can exacerbate the negative impact of ocean acidification on larvae and juveniles.     

Integrating effects of leaf nitrogen,age, rank,and growth temperature into the photosynthesis-stomatal conductance model LEAFC3-N parameterised for barley (Hordeum vulgare L.)

A crucial challenge for including biophysical photosynthesis–transpiration models into complex crop growth models is to integrate the plasticity of photosynthetic processes that is related to factors like nitrogen (N) content, age, and rank of leaves, or to the adaptation of plants to growth temperature (T_g). Here we present a new version of the combined photosynthesis-stomatal conductance model LEAFC3-N [Müller, J., Wernecke, P., Diepenbrock, W., 2005. LEAFC3-N: a nitrogen sensitive extension of the CO₂ and H₂O gas exchange model LEAFC3 parameterised and tested for winter wheat (Triticum aestivum L.). Ecological Modelling 183, 183–210.] that was revised, extended and completely re-parameterised for barley (Hordeum vulgare L.) with special regard for these factors to facilitate the use of the model in ecophysiological studies and in crop modelling. The analysis is based on novel comprehensive data on photosynthetic CO₂ and light response curves measured at two oxygen concentrations and different temperatures on leaves of barley (H. vulgare L.) differing in leaf N and chlorophyll content. Plants were grown in climatic chambers or in the field at different N and T_g.We thoroughly revised the existing and introduced new nitrogen relations for key model parameters that account for a linear increase with leaf N of V_max, J_max, T_p, and R_dmax (maximum rates of carboxylation, electron transport, triose phosphate export, and mitochondrial respiration), a saturation-type increase of φ (quantum yield of electron transport), and a non-linear decrease of θ and m (curvature of the light dependence of electron transport rate, scaling factor of the stomata model). The adaptation of photosynthetic characteristics to T_g was included into the model by linear relations that were observed between T_g and the activation energy ΔH_a of the temperature response characteristics of V_max, J_max, and T_p as well as of the nitrogen dependency of these characteristics. Based on an analysis of diurnal time courses of gas exchange rates it was found necessary including not only the relation between leaf water potential (Ψ) and stomatal conductance as used originally in LEAFC3, but additional effects on V_max and J_max. With the above-listed extensions, the model was capable to reproduce the observed plasticity and the recorded diurnal time courses of gas exchange rates fairly well. Thus, we conclude that the new model version can be used under a broad range of conditions, both for ecophysiological studies and as a submodel of crop growth models. The results presented here for barley will facilitate adapting photosynthesis models like LEAFC3-N to other C₃-species as well. The modelling of the effects of drought stress should be further elaborated in future based on more specific experiments.     

Effects of elevated pCO2 and the effect of parent acclimation on development in the tropical Pacific sea urchin Echinometra mathaei

Effects of acclimation to projected near-future ocean acidification (OA) conditions on physiology, reproduction and development were investigated in the tropical sea urchin Echinometra mathaei. Following 6 weeks in control or one of the three elevated pCO₂ (pH_NIST 7.5–8.1; pCO₂ ~485–1,770 μatm) conditions, adult urchins exhibited a slight decline of growth in low pH treatments and moderately reduced respiration at intermediate levels. At 7 weeks, gametes from adults were used to produce larvae that were reared in their respective parental treatments. To assess whether larvae from acclimated parents are more resilient to elevated pCO₂ than those not acclimated, larvae from control animals were also reared in the elevated pCO₂ treatments. There was no difference in female ‘spawnability’ and oocyte size between treatments, but male spawning ability was reduced in increased pCO₂ conditions. In elevated pCO₂ treatments, the percentage of normal larvae and larval size decreased in the progeny of control- and elevated pCO₂-acclimated parents, and arm asymmetry increased. Thus, acclimation of the parents did not make the progeny more resilient or sensitive to OA effects. Negative effects of increased pCO₂ on reproduction and development may impact on recruitment and population maintenance of this species.     

Application of a temperature-dependent von Bertalanffy growth model to bullhead (Cottus gobio)

The most studied and commonly applied model of fish growth is the von Bertalanffy model. However, this model does not take water temperature into account, which is one of the most important environmental factors affecting the life cycle of fish, as many physiological processes that determine growth, e.g. metabolic rate and oxygen supply, are directly influenced by temperature. In the present study we propose a version of the von Bertalanffy growth model that includes mean annual water temperatures by correlating the growth coefficient, k, explicitly and the asymptotic length, L_∞, implicitly to water temperature. All relationships include parameters with an obvious biological relevance that makes them easier to identify. The model is used to fit growth data of bullhead (Cottus gobio) at different locations in the Bez River network (Drme, France). We show that temperature explains much of the growth variability at the different sampling sites of the network.     

Effects of boreal forest management practices on the climate impact of CO2 emissions from bioenergy

In Life Cycle Assessment (LCA), carbon dioxide (CO₂) emissions from biomass combustion are traditionally assumed climate neutral if the bioenergy system is CO₂ flux neutral, i.e. the quantity of CO₂ released approximately equals the amount of CO₂ sequestered in biomass. This convention is a plausible assumption for fast growing biomass species, but is inappropriate for slower growing biomass, like forests. In this case, the climate impact from biomass combustion can be potentially underestimated if CO₂ emissions are ignored, or overestimated, if biogenic CO₂ is considered equal to anthropogenic CO₂. The estimation of the effective climate impact should take into account how the CO₂ fluxes are distributed over time: the emission of CO₂ from bioenergy approximately occurs at a single point in time, while the absorption by the new trees is spread over several decades. Our research target is to include this dynamic time dimension in unit-based impact analysis, using a boreal forest stand as case study. The boreal forest growth is modelled with an appropriate function, and is investigated under different forestry regimes (affecting the growth rate and the year of harvest). Specific atmospheric decay functions for biomass-derived CO₂ are then elaborated for selected combinations of forest management options. The contribution to global warming is finally quantified using the GWP_bio index as climate metric. Results estimates the effects of these practices on the characterization factor used for the global warming potential of CO₂ from bioenergy, and point out the key role played by the selected time horizon.     

Dynamic growth model for Scots pine (Pinus sylvestris L.) plantations in Galicia (north-western Spain)

In this study we developed a dynamic growth model for Scots pine (Pinus sylvestris L.) plantations in Galicia (north-western Spain). The data used to develop the model were obtained from a network of permanent plots, of between 10 and 55-year-old, which the Unidade de Xestión Forestal Sostible (Sustainable Forest Management Unit) of the University of Santiago de Compostela has set up in pure plantations of this species of pine in its area of distribution in Galicia. In this model, the initial stand conditions at any point in time are defined by three state variables (number of trees per hectare, stand basal area and dominant height), and are used to estimate stand volume, classified by commercial classes, for a given projection age. The model uses three transition functions expressed as algebraic difference equations of the three corresponding state variables used to project the stand state at any point in time. In addition, the model incorporates a function for predicting initial stand basal area, which can be used to establish the starting point for the simulation. This alternative should only be used when the stand is not yet established or when no inventory data are available. Once the state variables are known for a specific moment, a distribution function is used to estimate the number of trees in each diameter class, by recovering the parameters of the Weibull function, using the moments of first and second order of the distribution (arithmetic mean diameter and variance, respectively). By using a generalized height–diameter function to estimate the height of the average tree in each diameter class, combined with a taper function that uses the above predicted diameter and height, it is then possible to estimate total or merchantable stand volume.