In This Article Expand or collapse the "in this article" section Adaptive Landscapes

  • Introduction
  • General Overviews
  • Statistical Issues and Visualization of Fitness Surfaces
  • Role of Adaptive Landscapes in Molecular Evolution
  • Relationship between Adaptive Landscapes, Fitness Epistasis, and Linkage Disequilibrium
  • The Adaptive Landscape in Postzygotic Isolation and Speciation
  • Different Forms of Adaptive Landscapes
  • Landscape Movements on Microevolutionary Time Scales
  • Landscape Movements on Macroevolutionary Time Scales
  • Links to Other Fields and Role in Popular Science

Evolutionary Biology Adaptive Landscapes
Erik Svensson
  • LAST REVIEWED: 12 April 2023
  • LAST MODIFIED: 31 March 2016
  • DOI: 10.1093/obo/9780199941728-0073


Adaptive landscapes hold a central and special position in evolutionary theory, particularly in population and quantitative genetics, but also in some models of macroevolution. An adaptive landscape shows the relationship between fitness (vertical axis) and one or several traits or genes (horizontal axes). An adaptive landscape can therefore be viewed as a form of response surface, describing how a dependent variable (fitness) is causally influenced by one or several predictor variables (traits or genes). Evolution by natural selection in the context of an adaptive landscape can be viewed as a hill-climbing process, in which populations climb upwards to the trait or gene combination with the highest fitness, which are called “adaptive peaks.” In between the adaptive peaks, there are typically regions in phenotype or genotype space of low fitness (fitness valleys). Fitness valleys can also be thought of as the genetic combinations generating low-hybrid fitness when two incipient species meet and mate with each other, which generates a natural link between the adaptive landscape concept and speciation theory. In the original form and in the pre-computer era, adaptive landscapes were typically visualized as three-dimensional surface plots with only two traits (or genes). Adaptive landscapes did, in in their early days, mainly serve a heuristic function for qualitative reasoning, but over the last decades and with the development of evolutionary quantitative genetic theory and increasing computer power, they have now become a more quantitative tool for evolutionary biologists, including both theoreticians and empiricists. Whereas adaptive landscapes have become a part of mainstream evolutionary biology, they have also generated considerable controversy and confusion, as well as criticisms, mainly from philosophers. Some of these criticisms point to real conceptual problems, whereas others are based on misunderstandings of what the adaptive landscape actually is and what it can be used for.

General Overviews

The scientific literature on adaptive landscapes is huge, spanning both the philosophy and the evolutionary biology literature. This extensive literature includes journals, monographs, and edited books. A natural entry point for this literature is the population geneticist Sewall Wright’s original paper, “The Roles of Mutation, Inbreeding and Crossbreeding in Evolution” (Wright 1932). Frank and Slatkin 1992 discusses the different views of Sewall Wright and Ronald Fisher, arguing that Wright sought to establish a dynamical theory of evolution in his proposal of the adaptive landscape concept, whereas Fisher was more interested in evolutionary equilibria, through the formulation of his “fundamental theorem of natural selection” (Frank and Slatkin 1992). An excellent (although somewhat biased) monograph on Sewall Wright’s scientific career is provided in Provine 1986. Provine criticizes the adaptive landscape metaphor, and also points to inconsistencies between different versions of the adaptive landscape (the genotypic version versus the one on showing population allele frequencies), and suggests they are wholly incompatible. Dietrich and Skipper 2012 provides a brief history of the development of the adaptive landscape metaphor. Gavrilets 2004 discusses mainly genotypic fitness landscapes, starting with one- and two-locus models, and shows how such genotypic fitness landscapes can be used to model a wide range of phenomena in evolutionary biology, including the evolution of hybrid incompatibilities, sexual selection and sexual conflict, speciation, and molecular evolution. Gavrilets also shows how genotypic fitness landscapes and information about the fitnesses of individual genotypes can be used to deduce fitness landscapes at the population level (but notably, not the other way around), contra Provine’s claim. Barton and Turelli 1987 provides an early review of the role of adaptive landscapes in quantitative genetic theory. Arnold, et al. 2001 includes an extensive discussion about the differences between genotypic (Wrightian) adaptive landscapes and phenotypic (Simpsonian) adaptive landscapes, and reviews the quantitative genetic theory of how to quantify and measure adaptive peaks and valleys, building on the foundation set down by Lande 1976. Arnold and colleagues also argue that the adaptive landscape provides a natural conceptual bridge between micro- and macroevolution. Fear and Price 1998 discusses how the adaptive landscape can be used in investigations of ecology, and points out the difference between the individual fitness surface and the surface of population mean fitness. Schluter 2000 reviews the literature on adaptive radiation and ecological speciation theory, which is strongly based on the adaptive landscape tradition. Finally, the edited volume Svensson and Calsbeek 2012 contains a diversity of topics in chapters by historians, philosophers, evolutionary ecologists, and geneticists, who discuss the history, current scientific state, and future of the adaptive landscape from different perspectives.

  • Arnold, S. J., M. E. Pfrender, and A. G. Jones. 2001. The adaptive landscape as a conceptual bridge between micro- and macroevolution. Genetica 112–113:9–32.

    DOI: 10.1007/978-94-010-0585-2_2

    The authors argue that the scientific marriage of the “Simpsonian” adaptive landscape with evolutionary quantitative genetics is a natural framework for integrating micro- and macroevolutionary dynamics, including adaptive radiations, stabilizing selection, and peak shifts. They review mathematical modeling in the evolutionary quantitative genetics tradition, and argue that these models can shed lights on macroevolutionary processes.

  • Barton, N. H., and M. Turelli. 1987. Adaptive landscapes, genetic distance and the evolution of quantitative characters. Genetical Research 49:157–173.

    DOI: 10.1017/S0016672300026951

    An early and influential review of evolutionary quantitative genetics and its connection to adaptive landscape theory in evolutionary biology.

  • Dietrich, M. R., and R. A. Skipper. 2012. A shifting terrain: A brief history of the adaptive landscape. In The adaptive landscape in evolutionary biology. Edited by E. I. Svensson and R. Calsbeek, 3–15. Oxford: Oxford Univ. Press.

    An excellent historical review and introduction to how Sewall Wright came to formulate his idéa of the adaptive landscape.

  • Fear, K., and T. Price. 1998. The adaptive surface in ecology. Oikos 82:440–448.

    DOI: 10.2307/3546365

    The authors discuss how the adaptive landscape can be useful and informative to understand problems in ecology. They emphasize the crucial difference between mean fitness surface of the population and the individual fitness surface.

  • Frank, S. A., and M. Slatkin. 1992. Fisher’s fundamental theorem of natural selection. Trends in Ecology and Evolution 7:92–95.

    DOI: 10.1016/0169-5347(92)90248-A

    Explains the crucial difference between the two great population geneticists, Ronald Fisher and Sewall Wright, and their different views and interests. An excellent introduction to those who wish to understand the difference between “Fisherian” and “Wrightian” schools in population genetics and evolutionary biology.

  • Gavrilets, S. 2004. Fitness landscapes and the origin of species. Princeton, NJ: Princeton Univ. Press.

    In this influential monograph by a leading theoretical evolutionary biologist, Gavrilets show how genotypic fitness landscapes and information about the fitnesses of individual genotypes can be used to deduce fitness landscapes at the population level (but, notably, not the other way around), contra Provine’s claim (see Provine 1986).

  • Lande, R. 1976. Natural selection and random genetic drift in phenotypic evolution. Evolution 30:314–334.

    DOI: 10.2307/2407703

    A highly influential paper that formed the foundation for evolutionary quantitative genetics and established one of the first conceptual links between macroevolutionary processes as studied in the fossil record by paleontologists and what we know about the genetic basis of quantitative traits.

  • Provine, W. B. 1986. Sewall Wright and evolutionary biology. Chicago: Univ. of Chicago Press.

    A scholarly and authoritative biography on Sewall Wright, his scientific career, and his controversies with Fisher. Contains a critical, although somewhat subjective view of the adaptive landscape, which Provine is very critical of.

  • Schluter, D. 2000. The ecology of adaptive radiation. Oxford: Oxford Univ. Press.

    An influential and highly cited volume about the ecological causes of adaptive radiations and ecological speciation. Schluter’s analytical framework is largely built upon the adaptive landscape tradition.

  • Svensson, E. I., and R. E. Calsbeek, eds. 2012. The adaptive landscape in evolutionary biology. Oxford: Oxford Univ. Press.

    This edited volume also contains chapters dealing with how the adaptive landscape has influenced scientific research in animal behavior, conservation, development, micro- and macroevolution, paleontology, and speciation. Contributing authors come from different fields and include philosophers, historians of science, ecologists, geneticists, developmental and molecular biologists, and theoretical biologists.

  • Wright, S. 1932. The roles of mutation, inbreeding, crossbreeding and selection in evolution. In Proceedings of the Sixth International Congress of Genetics. Vol. 1, Transactions and general addresses. Edited by Donald F. Jones, 356–366. Austin, TX: Genetics Society of America.

    Sewall Wright’s classic paper from 1932, in which he outlined both his shifting balance theory (SBT) of evolution and the adaptive landscape concept, both of which have been very influential on evolutionary biology, but which have also caused considerable controversy.

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