Equilibrium and non-equilibrium concepts in forest genetic modelling: population- and individually-based approaches

Koen Kramer, D. C. van der Werf


The environment is changing and so are forests, in their functioning, in species composition, and in the species’ genetic composition. Many empirical and process-based models exist to support forest management. However, most of these models do not consider the impact of environmental changes and forest management on genetic diversity nor on the rate of adaptation of critical plant processes. How genetic diversity and rates of adaptation depend on management actions is a crucial next step in model development. Modelling approaches of genetic and demographic processes that operate in forests are categorized here in two classes. One approach assumes equilibrium conditions in phenotype and tree density, and analyses the characteristics of the demography and the genetic system of the species that determine the rate at which that equilibrium is attained. The other modelling approach does not assume equilibrium conditions and describes both the ecological —and genetic processes to analyse how environmental changes result in selection pressures on functional traits of trees and the consequences of that selection for tree— and ecosystem functioning. The equilibrium approach allows analysing the recovery rate after a perturbation in stable environments, i.e. towards the same pre-perturbation stable state. The nonequilibrium approach allows, in addition to the equilibrium approach, analysing consequences of ongoing environmental changes and forest management, i.e. non-stationary environments, on tree functioning, species composition, and genetic composition of the trees in forest ecosystem. In this paper we describe these two modelling approaches and discuss advantages and disadvantages of them and current knowledge gaps.


forest genetic models, population-genetic, individually-based models, equilibrium, non-equilibrium, environmental change, METAPOP, ForGEM

Full Text:



Austerlitz F., Mariette S., Machon N., Gouyon P.-H., Godelle B., 2000. Effects of colonization processes on genetic diversity: differences between annual plants and tree species. Genetics 1309-1321.
PMid:10757772 PMCid:1461003

Austin M.P., 1999. A silent clash of paradigms: some inconsistencies in community ecology. Oikos 86, 170-178.

Bouten W., 1995. Soil water dynamics of the solling spruce stand, calculated with the FORHYD simulation package. Ecological Modelling 83, 67-75.

Boyce M.S., 1998. Ecological-process management and ungulates: Yellowstone's conservation paradigm. Wildlife Society Bulletin 26, 391-398.

Briske D.D., Fuhlendorf S.D., Smeins F.E., 2003. Vegetation dynamics on rangelands: a critique of the current paradigm. J Appl Ecol 40, 601-614.

Champ J.G., Williams D.R., Knotek K., 2009. Wildland fire and organic discourse: negotiating place and leisure identity in a changing wildland urban interface. Leisure Sciences: An Interdisciplinary Journal, 31 237-254.

Corre V.L., Machon N., Petit R.J., Kremer A., 1997. Colonization with long-distance seed dispersal and genetic structure of maternally inherited genes in forest trees: a simulation study. Genet Res Camb 66, 117-125.

Deangelis D.L., Gross L.J., 1992. Individual-based models and approaches in ecology: populations, communities and ecosystems. Chapman & Hall Publishers 525 pp.

Dietze M.C., Clark J.S., 2008. Changing the gap dynamics paradigm: vegetative regeneration control on forest response to disturbance. Ecological Monographs 78, 331-347.

Falconer D.S., Mackay, T.F.C., 1996. Introduction to quantitative genetics. Addison Wesley Longman Ldt, Edinbrough. 464 pp.

Falconer D.S., Mackay T.F.C., 1996. Introduction to quantitative genetics, 4th ed. Adison Wesley Longman Group Ltd.

Geburek T., 2000. Effects of environmental pollution on the genetics of forest trees. In: Forest conservation genetics. Principles and practice (Young A., Boshier D., Boyle T., eds). CABI, Oxon, United Kindom.

Geburek T., Turok J., 2005. Conservation and management of forest genetic resources in Europe. Arbora Publishers, Zvolen.

Gladwin T.N., Kennelly J.J., Krause T.-S., 1995. Shifting paradigms for sustainable development: implications for management theory and research. Academy of Management Review 20, 874-907.

Gracia C., Fontes L., Bontemps J.-.D. et al., 2010. Models for forest management in a changing environment. Forest Systems.

Gregorius H.-R., 1978. The concept of genetic diversity and its formal relationship to heterozygozity and genetic distance. Math Biosc 41, 253-271.

Gregorius H.R., 1977. The genotype*environment-tophenotype relationship. Theoretical and Applied Genetics 49, 165-176.

Gregorius H.R., 1986. Measurements of genetical differentiation amoung subpopulations. Theoretical and Applied Genetics 71, 826-834.

Gregorius H.R., 1987. The relationship between the concepts of genetic diversity and differentiation. Theoretical and Applied Genetics 74, 397-401.

Gregorius H.R., 1988. The meaning of genetic variation within and between subpopulations. Theoretical and Applied Genetics 76, 947-951.

Hanssen A., 2002. Hessisches ministerium fuer umwelt, landwirtschaft und forsten. Hannover, Muenden. 142 pp.

Hartl D.L., Clark A.G., 1997. Principles of Population Genetics. Sinauer Associates, Inc., Sunderland, Massachusetts, 542 pp.

Hattemer H.H., 1991. Measuring genetic variation. In: Genetic variation in european populations of forest trees (Muller-Starck G.Z.M., ed). JD Sauerlander's Verlag, Frankfurt am Main. pp. 2-20.

Hengeveld R., 1997. Guest editorial: impact of biogeography on a population-biological paradigm shift. Journal of Biogeography 24, 541-547.

Hengeveld R., Walter G.H., 1999. The two coexisting ecological paradigms. Acta Biotheoretica 47, 141-170.

Huttunen S., 2009. Ecological modernisation and discourses on rural non-wood bioenergy production in Finland from 1980 to 2005. Journal of Rural Studies 25, 239-247.

Kotz S., Balakrishnan N., Johnson N.L., 2000. Continuous multivariate distributions. Models and applications. Wiley Press, Vol. 1. 752 pp.

Kramer K., 2007. In Climate change and forest genetic diversity: implications for sustainable forest management in Europe (Koskela J., Buck A., Teissier du Cros E., eds). IPGRI. Workdocument for Ministerial Conference on the Protection of Forests in Europe, April 2007, Poland, Paris, 15-16 March, 2006.

Kramer K., Brinkman A.G., Kuiters A.T., Verdonschot P.F.M., 2005. Alterra-report 1277. p. 80.

Kramer K., Buiteveld J., Forstreuter M. et al., 2008. Bridging the gap between ecophysiological and genetic knowledge to assess the adaptive potential of European beech. Ecological Modelling 216, 333-353.

Kramer K., Degen B., Buschbom J., Hickler T., Thuiller W., Sykes M.T., Winter W.D., 2010. Modelling exploration of the future of European beech (Fagus sylvatica L.) under climate change – Range, abundance, genetic diversity and adaptive response. Forest Ecology and Management 259, 2213-2222.

Kramer K., Baveco H., Bijlsma R.J.,Clerkx A.P.P.M., Dam J., Van Goethem J., Groen T.A., Groot Bruinderink G.W.T.A., Jorritsma I.T.M., Kalkhoven, Kuiters A.T.,Lammertsma D., Prins R.A., Sanders M., Wegman R., Van Wieren S.E., Wijdeven S., Van Der Wijngaart R., 2001. Alterra Report 216. Wageningen, The Netherlands. 168 pp.

Larsen A.B., 1996. Genetic structure of populations of beech (Fagus sylvatica L.) in Denmark. Scandinavian Journal of Forest Research 11, 220-232.

Le Corre V., Kremer A., 2003. Comparative dynamics of genetic variability of an adaptive trait and its underlying genes in a subdivided population. Genetics 1205-1219.
PMid:12871925 PMCid:1462636

Lomolino M.V., 2000. A call for a new paradigm of island biogeography. Global Ecology and Biogeography 9, 1-6.

Matala J., Karkkainen L., Harkonen K., Kellomaki S., Nuutinen T., 2009. Carbon sequestration in the growing stock of trees in Finland under different cutting and climate scenarios. European Journal of Forest Research, 128, 493-504.

Mátyás C., 1994. Modelling climate change effects with provenance test data. Tree Physiology, 14, 797-804.

Mátyás C., 1997. Perspectives of forest genetics and tree breeding in a changing world. Wien, 160 p pp.

Mitchell R.J., Duncan S.L., 2009. Range of variability in southern coastal plain forests: its historical, contemporary, and future role in sustaining biodiversity. Ecology and Society, 14, art.

O'Connor R.J., 2002. The conceptual basis of species distribution modelling: time for a paradigm shift? In: Predicting species occurrences: issues of accuracy and scale (Scott J.M., Heglund P.J., Haufler J., Morrison M., Raphael M., Wall B., Samson F., eds). Island Press, Covelo, California.

Rushton S.P., Ormerod S.J., Kerby G., 2004. New paradigms for modelling species distributions? J Appl Ecol 41, 193-200.

Simberloff D.S., 1980. A succession of paradigms in ecology: essentialism to materialism and probabilism. Synthese 43.

Solberg S., Dobbertin M., Reinds G.J. et al., 2009. Analyses of the impact of changes in atmospheric deposition and climate on forest growth in European monitoring plots: a stand growth approach. Forest Ecology and Management 258, 1735-1750.

Sunderlin W.D., 1995. Global environmental change, sociology, and paradigm isolation. Global Environmental Change Human and Policy Dimensions 5, 211-220.

Walter G.H., Hengeveld R., 2000. The structure of the two ecological paradigms. Acta Biotheoretica 48, 15-46.

Woo S.Y., 2009. Forest decline of the world: a linkage with air pollution and global warming. African Journal of Biotechnology 8, 7409-7414

Wu J., Loucks O.L., 1995. From balance of nature to hierarchical patch dynamics: a paradigm shift in ecology. Quarterly Review of Biology 70, 439-466.

DOI: 10.5424/fs/201019S-9312

Webpage: www.inia.es/Forestsystems