A parameterization of leaf phenology for the terrestrial ecosystem component of climate models

Leaf phenology remains one of the most difficult processes to parameterize in terrestrial ecosystem models because our understanding of the physical processes that initiate leaf onset and senescence is incomplete. While progress has been made at the molecular level, for example by identifying genes that are associated with senescence and flowering for selected plant species, a picture of the processes controlling leaf phenology is only beginning to emerge. A variety of empirical formulations have been used with varying degrees of success in terrestrial ecosystem models for both extratropical and tropical biomes. For instance, the use of growing degree-days (GDDs) to initiate leaf onset has received considerable recognition and this approach is used in a number of models. There are, however, limitations when using GDDs and other empirically based formulations in global transient climate change simulations. The phenology scheme developed for the Canadian Terrestrial Ecosystem Model (CTEM), designed for inclusion in the Canadian Centre for Climate Modelling and Analysis coupled general circulation model, is described. The representation of leaf phenology is general enough to be applied over the globe and sufficiently robust for use in transient climate change simulations. Leaf phenology is functionally related to the (possibly changing) climate state and to atmospheric composition rather than to geographical boundaries or controls implicitly based on current climate. In this approach, phenology is controlled by environmental conditions as they affect the carbon balance. A carbon-gain-based scheme initiates leaf onset when it is beneficial for the plant, in carbon terms, to produce new leaves. Leaf offset is initiated by unfavourable environmental conditions that incur carbon losses and these include shorter day length, cooler temperatures, and dry soil moisture conditions. The comparison of simulated leaf onset and offset times with observation-based estimates for temperate and boreal deciduous, tropical evergreen, and tropical deciduous plant functional types at selected locations indicates that the phenology scheme performs satisfactorily. Model simulated leaf area index and stem and root biomass are also compared with observational estimates to illustrate the performance of CTEM.

[1]  John L. Monteith,et al.  A reinterpretation of stomatal responses to humidity , 1995 .

[2]  R. Ahas,et al.  Changes in European spring phenology , 2002 .

[3]  A. P. Abaimov,et al.  Above- and belowground biomass and primary productivity of a Larix gmelinii stand near Tura, central Siberia. , 1999, Tree physiology.

[4]  R. Amasino,et al.  Vernalization in Arabidopsis thaliana is mediated by the PHD finger protein VIN3 , 2004, Nature.

[5]  Thomas Rötzer,et al.  Phenological maps of Europe , 2001 .

[6]  Andrea Molod,et al.  A New Look at Modeling Surface Heterogeneity: Extending Its Influence in the Vertical , 2003 .

[7]  T. A. Black,et al.  Diurnal and annual exchanges of mass and energy between an aspen‐hazelnut forest and the atmosphere: Testing the mathematical model Ecosys with data from the BOREAS experiment , 1999 .

[8]  J. Chen.  Optically-based methods for measuring seasonal variation of leaf area index in boreal conifer stands , 1996 .

[9]  M. V. van Wijk,et al.  Interannual variability of plant phenology in tussock tundra: modelling interactions of plant productivity, plant phenology, snowmelt and soil thaw , 2003 .

[10]  I. C. Prentice,et al.  An integrated biosphere model of land surface processes , 1996 .

[11]  Andrew White,et al.  Evaluation and analysis of a dynamic terrestrial ecosystem model under preindustrial conditions at the global scale , 2000 .

[12]  P. Warnant,et al.  CARAIB - A global model of terrestrial biological productivity , 1994 .

[13]  Hans Peter Schmid,et al.  Ecosystem‐atmosphere exchange of carbon dioxide over a mixed hardwood forest in northern lower Michigan , 2003 .

[14]  Wolfgang Knorr,et al.  Annual and interannual CO2 exchanges of the terrestrial biosphere: process-based simulations and uncertainties , 2000 .

[15]  I. C. Prentice,et al.  BIOME3: An equilibrium terrestrial biosphere model based on ecophysiological constraints, resource availability, and competition among plant functional types , 1996 .

[16]  R. Ceulemans,et al.  Stomatal conductance of forest species after long-term exposure to elevated CO2 concentration: a synthesis. , 2001, The New phytologist.

[17]  Peter M. Cox,et al.  Description of the "TRIFFID" Dynamic Global Vegetation Model , 2001 .

[18]  Takeshi Ohta,et al.  Climate change and extension of the Ginkgo biloba L. growing season in Japan , 2003 .

[19]  T. Black,et al.  Postulated feedbacks of deciduous forest phenology on seasonal climate patterns in the Western Canadian interior. , 2000 .

[20]  G. Whitelam,et al.  Gating of the rapid shade-avoidance response by the circadian clock in plants , 2003, Nature.

[21]  Vivek K. Arora,et al.  Simulating energy and carbon fluxes over winter wheat using coupled land surface and terrestrial ecosystem models , 2003 .

[22]  Jing M. Chen,et al.  Daily canopy photosynthesis model through temporal and spatial scaling for remote sensing applications , 1999 .

[23]  Vivek K. Arora,et al.  Streamflow simulations for continental-scale river basins in a global atmospheric general circulation model , 2001 .

[24]  Richard J. Norby,et al.  Phenological responses in maple to experimental atmospheric warming and CO2 enrichment , 2003 .

[25]  T. A. Black,et al.  Effects of seasonal and interannual climate variability on net ecosystem productivity of boreal deciduous and conifer forests , 2002 .

[26]  Mark D. Schwartz,et al.  Changes in North American spring , 2000 .

[27]  P. Reich,et al.  Extrapolating leaf CO2 exchange to the canopy: a generalized model of forest photosynthesis compared with measurements by eddy correlation , 1996, Oecologia.

[28]  J. Terborgh,et al.  The Phenology of Tropical Forests: Adaptive Significance and Consequences for Primary Consumers* , 1993 .

[29]  O. M. Heide.  High autumn temperature delays spring bud burst in boreal trees, counterbalancing the effect of climatic warming. , 2003, Tree physiology.

[30]  Matthias K.B. Lüdeke,et al.  The use of satellite NDVI data for the validation of global vegetation phenology models: application to the Frankfurt Biosphere Model , 1996 .

[31]  Michael T. Coe,et al.  Testing the performance of a dynamic global ecosystem model: Water balance, carbon balance, and vegetation structure , 2000 .

[32]  A. Menzel,et al.  Spatial and temporal variability of the phenological seasons in Germany from 1951 to 1996 , 2001 .

[33]  Olaf Kolle,et al.  Large carbon uptake by an unmanaged 250-year-old deciduous forest in Central Germany , 2002 .

[34]  Andrew D. Friend,et al.  A process-based, terrestrial biosphere model of ecosystem dynamics (Hybrid v3.0) , 1997 .

[35]  R. Grant.  Modeling topographic effects on net ecosystem productivity of boreal black spruce forests. , 2004, Tree physiology.

[36]  Xiaoqiu Chen,et al.  Relationships among phenological growing season, time‐integrated normalized difference vegetation index and climate forcing in the temperate region of eastern China , 2002 .

[37]  G. Kohlmaier,et al.  The Frankfurt Biosphere Model: a global process-oriented model of seasonal and long-term CO2 exchange between terrestrial ecosystems and the atmosphere. I. Model description and illustrative results for cold deciduous and boreal forests , 1994 .

[38]  C. Defila,et al.  Phytophenological trends in Switzerland , 2001, International journal of biometeorology.

[39]  Stephen P. Hubbell,et al.  Diameter, Height, Crown, and Age Relationship in Eight Neotropical Tree Species , 1995 .

[40]  J. Weber,et al.  5 – Environmental and Hormonal Control of Dormancy in Terminal Buds of Plants , 1978 .

[41]  A. Millar,et al.  How plants tell the time. , 2000, Current opinion in plant biology.

[42]  Thomas Rötzer,et al.  Annual and spatial variability of the beginning of growing season in Europe in relation to air temperature changes , 2002 .

[43]  R. Sage.  Acclimation of photosynthesis to increasing atmospheric CO2: The gas exchange perspective , 1994, Photosynthesis Research.

[44]  James M. Vose,et al.  Seasonal changes of leaf area index (LAI) in a tropical deciduous forest in west Mexico , 1995 .

[45]  H. Freeland,et al.  Spring phenology trends in Alberta, Canada: links to ocean temperature , 2000, International journal of biometeorology.

[46]  S. Yoshida.  Molecular regulation of leaf senescence. , 2003, Current opinion in plant biology.

[47]  R. Ahas.  Long-term phyto-, ornitho- and ichthyophenological time-series analyses in Estonia , 1999 .

[48]  Vivek K. Arora,et al.  A Representation of Variable Root Distribution in Dynamic Vegetation Models , 2003 .

[49]  R. Leuning.  A critical appraisal of a combined stomatal‐photosynthesis model for C3 plants , 1995 .

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

[51]  A. Henderson‐sellers,et al.  A global archive of land cover and soils data for use in general circulation climate models , 1985 .

[52]  D. Verseghy,et al.  Class—A Canadian land surface scheme for GCMS. I. Soil model , 2007 .

[53]  B. Bonan,et al.  A Land Surface Model (LSM Version 1.0) for Ecological, Hydrological, and Atmospheric Studies: Technical Description and User's Guide , 1996 .

[54]  Josep Peñuelas,et al.  Complex spatiotemporal phenological shifts as a response to rainfall changes. , 2004, The New phytologist.

[55]  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.

[56]  P. Reich,et al.  Relationships of leaf dark respiration to leaf nitrogen, specific leaf area and leaf life-span: a test across biomes and functional groups , 1998, Oecologia.

[57]  Annette Menzel,et al.  Growing season extended in Europe , 1999, Nature.

[58]  D. Verseghy,et al.  CLASS-A Canadian Land Surface Scheme for GCMs , 1993 .

[59]  J. Kummerow,et al.  Root biomass of a dry deciduous tropical forest in Mexico , 1991, Plant and Soil.

[60]  R. Norberg,et al.  Theory of Growth Geometry of Plants and Self-Thinning of Plant Populations: Geometric Similarity, Elastic Similarity, and Different Growth Modes of Plant Parts , 1988, The American Naturalist.

[61]  Eamus.  Ecophysiological traits of deciduous and evergreen woody species in the seasonally dry tropics. , 1999, Trends in ecology & evolution.

[62]  I. C. Prentice,et al.  Evaluation of ecosystem dynamics, plant geography and terrestrial carbon cycling in the LPJ dynamic global vegetation model , 2003 .

[63]  M. Jato,et al.  Pinus pollen in the atmosphere of Vigo and its relationship to meteorological factors , 2000, International journal of biometeorology.

[64]  T. Rötzer,et al.  Response of tree phenology to climate change across Europe , 2001 .

[65]  Primary Production by a Tropical Rain Forest of Southern Thailand , 1964 .

[66]  F. Berninger.  Effects of drought and phenology on GPP in Pinus sylvestris: a simulation study along a geographical gradient , 1997 .

[67]  The spatial distribution of forest biomass in the Brazilian Amazon: a comparison of estimates , 2001 .

[68]  J. Kauffman,et al.  Biomass, Carbon, and Nitrogen Pools in Mexican Tropical Dry Forest Landscapes , 2003, Ecosystems.

[69]  S. Running,et al.  A continental phenology model for monitoring vegetation responses to interannual climatic variability , 1997 .

[70]  Pierre Friedlingstein,et al.  A global prognostic scheme of leaf onset using satellite data , 2000 .

[71]  D. Randall,et al.  A Revised Land Surface Parameterization (SiB2) for Atmospheric GCMS. Part I: Model Formulation , 1996 .

[72]  Ann Henderson-Sellers,et al.  Biosphere-atmosphere transfer scheme(BATS) version 1e as coupled to the NCAR community climate model , 1993 .

[73]  S. Running,et al.  8 – Generalization of a Forest Ecosystem Process Model for Other Biomes, BIOME-BGC, and an Application for Global-Scale Models , 1993 .

[74]  Tiina Markkanen,et al.  Air temperature triggers the recovery of evergreen boreal forest photosynthesis in spring , 2003 .

[75]  Pierre Defourny,et al.  Seasonal variation of tropical forest LAI based on field measurements in Central African Republic , 2003 .

[76]  P. Friedlingstein,et al.  Toward an allocation scheme for global terrestrial carbon models , 1999 .