Capturing the superorganism: a formal theory of group adaptation

@article{Gardner2009CapturingTS,
  title={Capturing the superorganism: a formal theory of group adaptation},
  author={Andy Gardner and Alan Grafen},
  journal={Journal of Evolutionary Biology},
  year={2009},
  volume={22}
}
Adaptation is conventionally regarded as occurring at the level of the individual organism. However, in recent years there has been a revival of interest in the possibility for group adaptations and superorganisms. Here, we provide the first formal theory of group adaptation. In particular: (1) we clarify the distinction between group selection and group adaptation, framing the former in terms of gene frequency change and the latter in terms of optimization; (2) we capture the superorganism in… 
Group adaptation, formal darwinism and contextual analysis
TLDR
It is concluded that only clonal groups, and to a lesser extent groups in which reproductive competition is repressed, can be considered as adaptive units and an important distinction is focused on between two ways of understanding the links, which have different implications regarding group adaptationism.
Title: Group Adaptation, Formal Darwinism and Contextual Analysis
TLDR
It is concluded that only clonal groups, and to a lesser extent groups in which reproduc7 tive competition is repressed, can be considered as adaptive units.
A formal theory of the selfish gene
TLDR
A formal theory of the selfish gene is developed, using optimization theory to capture the analogy of ‘gene as fitness‐maximizing agent’ in mathematical terms and to assess the various criticisms that have been levelled at the theory, dispelling some and strengthening others.
The cultural evolution of emergent group-level traits
TLDR
The emergence and evolution of group-level traits and the implications for the theory of cultural evolution are discussed, including ramifications for the evolution of human cooperation, technology, and cultural institutions, and for the equivalency of multilevel selection and inclusive fitness approaches.
Group Selection and Group Adaptation During a Major Evolutionary Transition: Insights from the Evolution of Multicellularity in the Volvocine Algae
TLDR
It is suggested that group-specific adaptations could be present in the simplest, earliest branching colonial volvocine species, which do not have distinct specialized cells.
Levels of selection and the formal Darwinism project
TLDR
It is argued that the formal Darwinism project offers contradictory and confusing lines of thinking concerning level(s) of selection and suggested that a more even-handed approach, in which multicellular organisms are not assumed to have special status, would be desirable for a project that aims to make progress on the foundations of evolutionary theory.
Multilevel and kin selection in a connected world
TLDR
It is argued that the evolution of reduced virulence can be understood from the perspective of inclusive fitness, obviating the need to evoke group selection as a contributing causal factor, and that the cause of reducedvirulence resides in the opposition of two processes: within-group and among-group selection.
A levels-of-selection approach to evolutionary individuality
TLDR
A method by which the property ‘evolutionary individuality’ can be measured is suggested and it is argued that a problem which is often considered to be fatal to that method—the problem of ‘cross-level by-products’—can be avoided.
The genetical theory of multilevel selection
  • A. Gardner
  • Psychology
    Journal of evolutionary biology
  • 2015
TLDR
It is shown that taking a genetical approach facilitates a decomposition of group‐level traits – including reproductive success – into the separate contributions made by each constituent individual, even in the context of so‐called emergence.
The evolution of cooperation, especially in humans
TLDR
It is shown that in the simplest case, because of the cost of competition between non-dispersing siblings, the level of promiscuity has little or no effect on the evolution of cooperation, and between-group competition can favour any social behaviour so long as it helps the group compete.
...
...

References

SHOWING 1-10 OF 113 REFERENCES
Reviving the superorganism.
ARE WE STALLED PART WAY THROUGH A MAJOR EVOLUTIONARY TRANSITION FROM INDIVIDUAL TO GROUP?
  • S. Stearns
  • Psychology
    Evolution; international journal of organic evolution
  • 2007
This commentary poses an evolutionary hypothesis about the nature of the human condition: that we are stalled part way through a major evolutionary transition from individuals to groups, a transition
Alternative formulations of multilevel selection
TLDR
It is argued that the “effect hypothesis” of macroevolution, requiring, in selection among species, both the absence of group effects on organismic fitness (multilevel selection) and the direct determination of species fitnesses by those of organisms, is untestable with paleontological data.
Eusociality: origin and consequences.
In this new assessment of the empirical evidence, an alternative to the standard model is proposed: group selection is the strong binding force in eusocial evolution; individual selection, the strong
The emergence of a superorganism through intergroup competition
Surveys of insect societies have revealed four key, recurring organizational trends: (i) The most elaborated cooperation occurs in groups of relatives. (ii) Cooperation is typically more elaborate in
The formal Darwinism project: a mid‐term report
  • A. Grafen
  • Psychology
    Journal of evolutionary biology
  • 2007
TLDR
The ‘Formal Darwinism Project’ is pursued, whose essence is to underpin and formalize the fitness optimization ideas used by behavioural ecologists, using a new kind of argument linking the Mathematics of motion and the mathematics of optimization.
Evolution of individuality during the transition from unicellular to multicellular life
  • R. Michod
  • Biology
    Proceedings of the National Academy of Sciences
  • 2007
TLDR
The hypothesis is that fitness tradeoffs drive the transition of a cell group into a multicellular individual through the evolution of cells specialized at reproductive and vegetative functions of the group.
Rethinking the Theoretical Foundation of Sociobiology
TLDR
This article takes a “back to basics” approach, explaining what group selection is, why its rejection was regarded as so important, and how it has been revived based on a more careful formulation and subsequent research.
The genetical evolution of social behaviour. I.
A Method for Analyzing Selection in Hierarchically Structured Populations
Individual fitness depends on the particular ecological, genetic, and social contexts in which organisms are found. Variation in individual context among subunits of a population thus raises
...
...