Contributions of climate, leaf area index and leaf physiology to variation in gross primary production of six coniferous forests across Europe: a model-based analysis.

Gross primary production (GPP) is the primary source of all carbon fluxes in the ecosystem. Understanding variation in this flux is vital to understanding variation in the carbon sink of forest ecosystems, and this would serve as input to forest production models. Using GPP derived from eddy-covariance (EC) measurements, it is now possible to determine the most important factor to scale GPP across sites. We use long-term EC measurements for six coniferous forest stands in Europe, for a total of 25 site-years, located on a gradient between southern France and northern Finland. Eddy-derived GPP varied threefold across the six sites, peak ecosystem leaf area index (LAI) (all-sided) varied from 4 to 22 m(2) m(-2) and mean annual temperature varied from -1 to 13 degrees C. A process-based model operating at a half-hourly time-step was parameterized with available information for each site, and explained 71-96% in variation between daily totals of GPP within site-years and 62% of annual total GPP across site-years. Using the parameterized model, we performed two simulation experiments: weather datasets were interchanged between sites, so that the model was used to predict GPP at some site using data from either a different year or a different site. The resulting bias in GPP prediction was related to several aggregated weather variables and was found to be closely related to the change in the effective temperature sum or mean annual temperature. High R(2)s resulted even when using weather datasets from unrelated sites, providing a cautionary note on the interpretation of R(2) in model comparisons. A second experiment interchanged stand-structure information between sites, and the resulting bias was strongly related to the difference in LAI, or the difference in integrated absorbed light. Across the six sites, variation in mean annual temperature had more effect on simulated GPP than the variation in LAI, but both were important determinants of GPP. A sensitivity analysis of leaf physiology parameters showed that the quantum yield was the most influential parameter on annual GPP, followed by a parameter controlling the seasonality of photosynthesis and photosynthetic capacity. Overall, the results are promising for the development of a parsimonious model of GPP.

[1]  G. Grassi,et al.  Foliar morphological and physiological plasticity in Picea abies and Abies alba saplings along a natural light gradient. , 2001, Tree physiology.

[2]  D. Loustau,et al.  Variability of the photosynthetic characteristics of mature needles within the crown of a 25-year-old Pinus pinaster. , 1998, Tree physiology.

[3]  T. Vesala,et al.  Deriving a light use efficiency model from eddy covariance flux data for predicting daily gross primary production across biomes , 2007 .

[4]  Pauline Stenberg,et al.  Simulations of the effects of shoot structure and orientation on vertical gradients in intercepted light by conifer canopies. , 1996, Tree physiology.

[5]  S. Running,et al.  A general model of forest ecosystem processes for regional applications I. Hydrologic balance, canopy gas exchange and primary production processes , 1988 .

[6]  Dennis D. Baldocchi,et al.  On using eco-physiological, micrometeorological and biogeochemical theory to evaluate carbon dioxide, water vapor and trace gas fluxes over vegetation: a perspective , 1998 .

[7]  Tiina Markkanen,et al.  Effect of thinning on surface fluxes in a boreal forest , 2005 .

[8]  C. Field,et al.  Scaling physiological processes: leaf to globe. , 1995 .

[9]  Eero Nikinmaa,et al.  Developing an empirical model of stand GPP with the LUE approach: analysis of eddy covariance data at five contrasting conifer sites in Europe , 2007 .

[10]  W. Oechel,et al.  A new model of gross primary productivity for North American ecosystems based solely on the enhanced vegetation index and land surface temperature from MODIS , 2008 .

[11]  A. Mäkelä,et al.  Acclimation of photosynthetic capacity in Scots pine to the annual cycle of temperature. , 2004, Tree physiology.

[12]  I. Mammarella,et al.  Determining the contribution of vertical advection to the net ecosystem exchange at Hyytiälä forest, Finland , 2007 .

[13]  M. Adams,et al.  Photosynthesis-Rubisco relationships in foliage of Pinus sylvestris in response to nitrogen supply and the proposed role of Rubisco and amino acids as nitrogen stores , 2003, Trees.

[14]  T. Pukkala,et al.  Relationship between radiation interception and photosynthesis in forest canopies: effect of stand structure and latitude , 1989 .

[15]  A. J. Dolman,et al.  Radiation, temperature, and leaf area explain ecosystem carbon fluxes in boreal and temperate European forests , 2005 .

[16]  P. Berbigier,et al.  CO2 and water vapour fluxes for 2 years above Euroflux forest site , 2001 .

[17]  R. Ceulemans,et al.  Scaling-Up Carbon Fluxes from Leaves to Stands in a Patchy Coniferous / Deciduous Forest , 1997 .

[18]  G. Campbell,et al.  An Introduction to Environmental Biophysics , 1977 .

[19]  P. De Angelis,et al.  Effects of elevated (CO2) on photosynthesis in European forest species: a meta-analysis of model parameters , 1999 .

[20]  Frank Veroustraete,et al.  Seasonal variations in leaf area index, leaf chlorophyll, and water content; scaling-up to estimate fAPAR and carbon balance in a multilayer, multispecies temperate forest. , 1999, Tree physiology.

[21]  Philip J. Radtke,et al.  Bayesian melding of a forest ecosystem model with correlated inputs , 2002 .

[22]  Sean C. Thomas,et al.  The worldwide leaf economics spectrum , 2004, Nature.

[23]  Markus Reichstein,et al.  Evidence for soil water control on carbon and water dynamics in European forests during the extremely dry year: 2003 , 2007 .

[24]  Robert Clement,et al.  A comparative analysis of simulated and observed photosynthetic CO2 uptake in two coniferous forest canopies. , 2006, Tree physiology.

[25]  Henry L. Gholz,et al.  Environmental Limits on Aboveground Net Primary Production, Leaf Area, and Biomass in Vegetation Zones of the Pacific Northwest , 1982 .

[26]  R. Waring,et al.  A generalised model of forest productivity using simplified concepts of radiation-use efficiency, carbon balance and partitioning , 1997 .

[27]  P. Hari,et al.  Instantaneous PAR estimated using long records of daily temperature and rainfall , 2001 .

[28]  S. Running,et al.  Leaf Area of Mature Northwestern Coniferous Forests: Relation to Site Water Balance , 1977 .

[29]  D. Baldocchi,et al.  CO2 fluxes over plant canopies and solar radiation: a review , 1995 .

[30]  Belinda E. Medlyn,et al.  Energy Conversion and Use in Forests: An Analysis of Forest Production in Terms of Radiation Utilisation Efficiency (ɛ) , 1997 .

[31]  T. Vesala,et al.  Simulation and scaling of temporal variation in gross primary production for coniferous and deciduous temperate forests , 2004 .

[32]  Tiina Markkanen,et al.  Interannual variability and timing of growing‐season CO2 exchange in a boreal forest , 2003 .

[33]  Paul G. Jarvis,et al.  Description and validation of an array model - MAESTRO. , 1990 .

[34]  S. T. Gower,et al.  A cross‐biome comparison of daily light use efficiency for gross primary production , 2003 .

[35]  W. Oechel,et al.  Seasonality of ecosystem respiration and gross primary production as derived from FLUXNET measurements , 2001 .

[36]  R. Ceulemans,et al.  Footprint-adjusted net ecosystem CO2 exchange and carbon balance components of a temperate forest , 2006 .

[37]  P. Hari,et al.  Comparison of an optimal stomatal regulation model and a biochemical model in explaining CO‚́‚ exchange in field conditions , 2002 .

[38]  I. R. Cowan,et al.  Stomatal function in relation to leaf metabolism and environment. , 1977, Symposia of the Society for Experimental Biology.

[39]  U. Niinemets Stomatal conductance alone does not explain the decline in foliar photosynthetic rates with increasing tree age and size in Picea abies and Pinus sylvestris. , 2002, Tree physiology.

[40]  Arnaud Carrara,et al.  Net ecosystem CO2 exchange of mixed forest in Belgium over 5 years , 2003 .

[41]  I. R. Cowan,et al.  INTEGRATION OF ACTIVITY IN THE HIGHER PLANT , 1977 .

[42]  F. Meinzer,et al.  Boundary layer conductance, leaf temperature and transpiration of Abies amabilis branches. , 1999, Tree physiology.

[43]  D. A. Sampson,et al.  Basal rates of soil respiration are correlated with photosynthesis in a mixed temperate forest , 2007 .

[44]  Robert Clement,et al.  On the validation of models of forest CO2 exchange using eddy covariance data: some perils and pitfalls. , 2005, Tree physiology.

[45]  M. Aurela Carbon dioxide exchange in subarctic ecosystems measured by a micrometeorological technique , 2005 .

[46]  Zisheng Xing,et al.  Total and component carbon fluxes of a Scots pine ecosystem from chamber measurements and eddy covariance. , 2007, Annals of botany.

[47]  F. Berninger,et al.  Modeling 13C discrimination in tree rings , 2000 .

[48]  A. Mäkelä,et al.  Optimal control of gas exchange. , 1986, Tree physiology.

[49]  Ü. Rannik,et al.  Gap filling strategies for defensible annual sums of net ecosystem exchange , 2001 .

[50]  John M. Norman,et al.  Carbon distribution and aboveground net primary production in aspen, jack pine, and black spruce stands in Saskatchewan and Manitoba, Canada , 1997 .

[51]  K. Hibbard,et al.  A Global Terrestrial Monitoring Network Integrating Tower Fluxes, Flask Sampling, Ecosystem Modeling and EOS Satellite Data , 1999 .

[52]  Lars Eklundh,et al.  Net primary production and light use efficiency in a mixed coniferous forest in Sweden , 2005 .

[53]  Alan R. Ek,et al.  Process-based models for forest ecosystem management: current state of the art and challenges for practical implementation. , 2000, Tree physiology.

[54]  E. Falge,et al.  Effects of stand structure and physiology on forest gas exchange: a simulation study for Norway spruce , 1997, Trees.

[55]  J. Berry,et al.  A biochemical model of photosynthetic CO2 assimilation in leaves of C3 species , 1980, Planta.

[56]  A. Mäkelä,et al.  Annual pattern of photosynthesis in Scots pine in the boreal zone. , 2003, Tree physiology.

[57]  B. Medlyn,et al.  Temperature response of parameters of a biochemically based model of photosynthesis. I. Seasonal changes in mature maritime pine (Pinus pinaster Ait.) , 2002 .

[58]  Christian Bernhofer,et al.  A decade of carbon, water and energy flux measurements of an old spruce forest at the Anchor Station Tharandt , 2007 .

[59]  J. William Munger,et al.  Exchange of Carbon Dioxide by a Deciduous Forest: Response to Interannual Climate Variability , 1996, Science.

[60]  A. Mäkelä,et al.  Field evidence for the optimality hypothesis of gas exchange in plants , 1999 .

[61]  G. Mohren,et al.  Impacts of Global Change on Tree Physiology and Forest Ecosystems , 1997, Forestry Sciences.

[62]  P. Hari,et al.  Long-term field measurements of atmosphere-surface interactions in boreal forest combining forest ecology, micrometeorology, aerosol physics and atmospheric chemistry , 1998 .

[63]  J. Landsberg,et al.  Evaluating a simple radiation/dry matter conversion model using data from Eucalyptus globulus plantations in Western Australia. , 1996, Tree physiology.

[64]  S. Linder,et al.  Photosynthesis and transpiration in 20-year-old Scots pine. , 1980 .

[65]  Denis Loustau,et al.  Carbon balance of coniferous forests growing in contrasting climates: model-based analysis , 2005 .

[66]  B. Guan Effects of correlation among parameters on prediction quality of a process-based forest growth model. , 2000 .

[67]  W. Oechel,et al.  Environmental controls over carbon dioxide and water vapor exchange of terrestrial vegetation , 2002 .

[68]  Tiina Markkanen,et al.  Eddy covariance fluxes over a boreal Scots pine forest , 2001 .

[69]  Markus Reichstein,et al.  Cross-site evaluation of eddy covariance GPP and RE decomposition techniques , 2008 .

[70]  John M. Norman,et al.  4 – Scaling Processes between Leaf and Canopy Levels , 1993 .

[71]  A. Mäkelä,et al.  Predicting the decline in daily maximum transpiration rate of two pine stands during drought based on constant minimum leaf water potential and plant hydraulic conductance. , 2008, Tree physiology.

[72]  Denis Loustau,et al.  Temperature response of parameters of a biochemically based model of photosynthesis. II. A review of experimental data , 2002 .

[73]  Heikki Hänninen,et al.  Relationship between temperature and the seasonal course of photosynthesis in Scots pine at northern timberline and in southern boreal zone , 2007 .

[74]  L. Kergoat A model for hydrological equilibrium of leaf area index on a global scale , 1998 .

[75]  C. Pye-Smith From the forest floor , 2005 .

[76]  E. Davidson,et al.  Comparing simple respiration models for eddy flux and dynamic chamber data , 2006 .

[77]  Eero Nikinmaa,et al.  Forest floor vegetation plays an important role in photosynthetic production of boreal forests , 2006 .

[78]  Sune Linder,et al.  Climatic factors controlling the productivity of Norway spruce : A model-based analysis , 1998 .

[79]  Ü. Rannik,et al.  Estimates of the annual net carbon and water exchange of forests: the EUROFLUX methodology , 2000 .

[80]  A. Mäkelä,et al.  Modelling five years of weather-driven variation of GPP in a boreal forest , 2006 .

[81]  D. Reichle,et al.  Dynamic properties of forest ecosystems , 2004, Vegetatio.

[82]  W. Cohen,et al.  Scaling Gross Primary Production (GPP) over boreal and deciduous forest landscapes in support of MODIS GPP product validation , 2003 .

[83]  W. S. Benninghoff Structure and Function of Northern Coniferous Forests—An Ecosystem Study , 1984 .

[84]  S. Luoma Geographical pattern in photosynthetic light response of Pinus sylvestris in Europe , 1997 .

[85]  John Philip Cooper,et al.  Photosynthesis and productivity in different environments , 1976 .

[86]  Continuous long-term measurements of soil-plant-atmosphere variables at a forest site , 1999 .

[87]  T. Vesala,et al.  On the separation of net ecosystem exchange into assimilation and ecosystem respiration: review and improved algorithm , 2005 .

[88]  M. Aubinet,et al.  Determinants of terrestrial ecosystem carbon balance inferred from European eddy covariance flux sites , 2007 .

[89]  Henry L. Gholz,et al.  The Use of Remote Sensing in the Modeling of Forest Productivity , 1997, Forestry Sciences.

[90]  D. Baldocchi,et al.  The human footprint in the carbon cycle of temperate and boreal forests − , 2008 .

[91]  S. Running,et al.  The impact of growing-season length variability on carbon assimilation and evapotranspiration over 88 years in the eastern US deciduous forest , 1999, International journal of biometeorology.

[92]  A. Sellin Morphological and stomatal responses of Norway spruce foliage to irradiance within a canopy depending on shoot age. , 2001, Environmental and experimental botany.