title: Adaptive multiscapes: an up-to-date metaphor to visualize molecular adaptation.
abstract: Wright's metaphor of the fitness landscape has shaped and conditioned our view of the adaptation of populations for almost a century. Since its inception, and including criticism raised by Wright himself, the concept has been surrounded by controversy. Among others, the debate stems from the intrinsic difficulty to capture important features of the space of genotypes, such as its high dimensionality or the existence of abundant ridges, in a visually appealing two-dimensional picture. Two additional currently widespread observations come to further constrain the applicability of the original metaphor: the very skewed distribution of phenotype sizes (which may actively prevent, due to entropic effects, the achievement of fitness maxima), and functional promiscuity (i.e. the existence of secondary functions which entail partial adaptation to environments never encountered before by the population).
  
  
  
  Here we revise some of the shortcomings of the fitness landscape metaphor and propose a new "scape" formed by interconnected layers, each layer containing the phenotypes viable in a given environment. Different phenotypes within a layer are accessible through mutations with selective value, while neutral mutations cause displacements of populations within a phenotype. A different environment is represented as a separated layer, where phenotypes may have new fitness values, other phenotypes may be viable, and the same genotype may yield a different phenotype, representing genotypic promiscuity. This scenario explicitly includes the many-to-many structure of the genotype-to-phenotype map. A number of empirical observations regarding the adaptation of populations in the light of adaptive multiscapes are reviewed.
  
  
  
  Several shortcomings of Wright's visualization of fitness landscapes can be overcome through adaptive multiscapes. Relevant aspects of population adaptation, such as neutral drift, functional promiscuity or environment-dependent fitness, as well as entropic trapping and the concomitant impossibility to reach fitness peaks are visualized at once. Adaptive multiscapes should aid in the qualitative understanding of the multiple pathways involved in evolutionary dynamics.
  
  
  
  This article was reviewed by Eugene Koonin and Ricard Solé.
authors.affiliation: Grupo Interdisciplinar de Sistemas Complejos (GISC), Madrid, Spain.|Departamento de Matemáticas, Universidad Carlos III de Madrid, Madrid, Spain.
authors.forename: Pablo
authors.initials: P
authors.lastname: Catalán
authors.affiliation: Grupo Interdisciplinar de Sistemas Complejos (GISC), Madrid, Spain.
authors.forename: Clemente F
authors.initials: CF
authors.lastname: Arias
authors.affiliation: Grupo Interdisciplinar de Sistemas Complejos (GISC), Madrid, Spain.|Departamento de Matemáticas, Universidad Carlos III de Madrid, Madrid, Spain.|Institute for Biocomputation and Physics of Complex Systems, Zaragoza, Spain.|UC3M-BS Institute of Financial Big Data (IFiBiD), Madrid, Spain.
authors.forename: Jose A
authors.initials: JA
authors.lastname: Cuesta
authors.affiliation: Grupo Interdisciplinar de Sistemas Complejos (GISC), Madrid, Spain. smanrubia@cnb.csic.es.|National Biotechnology Centre (CSIC), c/ Darwin 3, Madrid, 28049, Spain. smanrubia@cnb.csic.es.
authors.forename: Susanna
authors.identifier: 0000-0003-0134-2785
authors.initials: S
authors.lastname: Manrubia
mesh_terms: D000220:Adaptation, Biological; D019143:Evolution, Molecular; D005838:Genotype; D008957:Models, Genetic; D009154:Mutation; D010641:Phenotype; D011157:Population Dynamics; D012641:Selection, Genetic
keywords: Adaptive landscape; Environment; Functional promiscuity; Genotype-phenotype map; Neutral networks; Phenotype size