Social selection

Last updated
Mutation and Selection Mutation and selection diagram (2).svg
Mutation and Selection

Social selection is a term used with varying meanings in biology.

Contents

Joan Roughgarden proposed a hypothesis called social selection as an alternative to sexual selection. Social selection is argued to be a mode of natural selection based on reproductive transactions and a two-tiered approach to evolution and the development of social behavior. [1] Reproductive transactions refer to a situation where one organism offers assistance to another in exchange for access to reproductive opportunity. The two tiers of the theory are behavioral and population genetic. [1] [2] The genetic aspect states that anisogamy arose to maximize contact rate between gametes. The behavioral aspect is concerned with cooperative game theory and the formation of social groups to maximize the production of offspring. In her critique against the neo-Darwinian defense of sexual selection, Roughgarden outlines exceptions to many of the assumptions that come with sexual selection. [1] These exceptions include sexually monomorphic species, species which reverse standard sex roles, species with template multiplicity, species with transgender presentation, frequencies of homosexual mating, and the lack of correlation between sexually selected traits and deleterious mutation. [3]

An article published by Roughgarden's lab on her ideas received criticism in the journal Science . Forty scientists produced ten critical letters. The critics stated that the article was misleading, that it contained misunderstandings and misrepresentations, that sexual selection accounted for all the data presented and subsumed Roughgarden's theoretical analysis, and that sexual selection explained data that her theory could not. [4] [5]

Other researchers, such as biologist Mary Jane West-Eberhard and evolutionary medicine researcher Randolph M. Nesse, instead view sexual selection as a subcategory of social selection, [list 1] with Nesse and anthropologist Christopher Boehm arguing further that altruism in humans held fitness advantages that enabled evolutionarily extraordinary cooperativeness and the human capability of creating culture, as well as capital punishment by band societies against bullies, thieves, free-riders, and psychopaths. [list 2]

Genetic principles

Portfolio hypothesis

Short for the genetic-portfolio balancing hypothesis, this idea, proposed by Roughgarden, is used as an alternative to the Red Queen and Mueller's ratchet hypotheses to explain the existence of sexual reproduction within the framework of social selection. [1] In a population with two species which fit the same ecological niche, live in the same local environment, have the same degree of genetic diversity, but have different modes of reproduction, sexual and asexual, the sexual species will eventually dominate the local environment. This is due to asexual populations losing diversity for short-term adaptations to the environment.

Roughgarden proposes a population of dandelions which fit the above description. The parental generation of a sexually reproducing species and asexually reproducing species contains equal ratios of the three genotypes (A1A1, A1A2, and A2A2). The F1 generation of the asexual dandelions will contain the same ratio as the P-generation. Conversely, following standard principles of sexual reproduction, the F1 generation will be 25% A1A1, 25% A2A2, and 50% A1A2. With the addition of differential survival related to these genotypes (certain genotypes surviving better in different degrees of sunlight), the asexual population will eventually drift toward one genotype and die off when the environment changes to suit a different genotype. The sexual population in the same situation will remain diverse enough to survive changing environments.

From this theory, Roughgarden concludes that the main benefit of sexual reproduction is the maintenance of genetic diversity when compared to similar asexual populations.

The IR model of the development of anisogamy

The IR model for the development of anisogamy is named after its developers Priya Iyer and Joan Roughgarden. By considering the evolution of anisogamy in hermaphroditic marine invertebrates and bisexual plants, the theory postulates of a gene locus which controls both sperm and egg size produced by an organism. Anisogamy could evolve in diploid hermaphroditic adults as an individual adaptation which increases its own fitness. [18]

Hermaphrodism

Hermaphroditic animals and dioecious plants represent a large portion of sexually reproductive species. Under social selection theory, species where individuals produce two different gametes predate strictly gonochoristic and monoecious species. Separate sexes can, therefore, be described as derivations of primal hermaphrodites. [1]

Males arising in primarily hermaphroditic species gain an advantage in certain environments as fertilizers because they lack the energy cost of producing eggs. [19] The development of monoecious and gonochoristic species represents a transition from broadcast fertilization to localized and internal fertilization. [1]

The Broad-barred goby is capable of bi-directional sex change. Gobiodon histrio 6.1.2011 030 b.jpg
The Broad-barred goby is capable of bi-directional sex change.

Simultaneous hermaphrodism exists in species with pre-Cambrian roots, and several families of organisms have shifted between hermaprodism and gonochoism over their evolutionary history. [1] There are sequentially hermaphroditic species, such as the goby, which show bidirectional sex changing. The dwarf males of anglerfish in the family Ceratiidae function as "mobile testes" for the females of their species.

Behavioral principles

Reproductive transactions

Animals help another in order to access reproductive opportunities. Any inequality in this opportunity is due to predation or resource availability. Therefore, there is value in boosting the reproductive fitness of an animal's parents or siblings, both of which share genetic information. Even without this genetic relationship, reproductive transactions can be valuable. Sexual conflict arises from a failure for pairs to negotiate value of reproductive transactions effectively. [1]

Mating can therefore serve purposes beyond reproduction, if the maintenance of social structures does not decrease effective fitness. Homosexual mating behavior is observed in species where this is the case. Several asexual species of whiptail lizards have been observed to engage in mating and pair-bonding despite the lack of gametic fusion. [20]

Social state matrices

Animal behavior can be understood as the intersection of three primary elements: genetic foundations, social systems, and individual reaction. A social state matrix is composed of genetic foundations and social systems to determine behavior arising from the intersection of them. [1] For example, animals have genetics which determine reaction to potential foraging stimuli, but only search for the stimuli at certain times of day due to social systems. Therefore, social systems would be selected for which optimize behaviors such as foraging and mate selection.

Criticism

Presented as an alternative to sexual selection theory, social selection has received criticism as a result. Arguments have been made that Roughgarden anthropomorphizes animal behavior in order to suit her theory. [21] Other critics argue that the holes in sexual selection theory which Roughgarden proposes, such as inconsistencies in male and female relationships and critiques of Bateman's principle, can actually be consolidated within the sexual selection framework. [22]

An article published by Roughgarden's lab on these ideas received criticism in the journal Science . Forty scientists produced ten critical letters. The critics stated that the article was misleading, that it contained misunderstandings and misrepresentations, that sexual selection accounted for all the data presented and subsumed Roughgarden's theoretical analysis, and that sexual selection explained data that her theory could not. [4] [5] Roughgarden stated she was "not altogether surprised" by the volume of dissent and that her theory was not an extension of sexual selection theory. [4] [5]

Alternate uses of the term

The term "social selection" has been used by other researchers to describe elements of the selection process overlooked by the theory of sexual selection, and to view sexual selection as a subcategory of social selection. [6] [7] [11] Mary Jane West-Eberhard used the term social selection to describe differential success in social competition for resources other than mates, [8] which includes female competition for territory and competition for parental attention among offspring. [9] Citing cross-cultural research conducted by social psychologist David Buss, [23] [24] psychologist Geoffrey Miller has argued that if humans prefer altruistic mating partners that would select by mate choice for altruism directly, [25] while evolutionary medicine researcher Randolph M. Nesse has argued that humans with altruistic tendencies receive fitness advantages because they are preferred as social partners, [14] and this enabled humans as a species of becoming extraordinarily cooperative and capable of creating culture. [15]

Related Research Articles

Evolutionary psychology is a theoretical approach in psychology that examines cognition and behavior from a modern evolutionary perspective. It seeks to identify human psychological adaptations with regards to the ancestral problems they evolved to solve. In this framework, psychological traits and mechanisms are either functional products of natural and sexual selection or non-adaptive by-products of other adaptive traits.

<span class="mw-page-title-main">Sexual selection</span> Mode of natural selection involving the choosing of and competition for mates

Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with, and compete with members of the same sex for access to members of the opposite sex. These two forms of selection mean that some individuals have greater reproductive success than others within a population, for example because they are more attractive or prefer more attractive partners to produce offspring. Successful males benefit from frequent mating and monopolizing access to one or more fertile females. Females can maximise the return on the energy they invest in reproduction by selecting and mating with the best males.

<span class="mw-page-title-main">Sex</span> Trait that determines an organisms sexually reproductive function

Sex is the trait that determines whether a sexually reproducing organism produces male or female gametes. During sexual reproduction, a male and a female gamete fuse to form a zygote, which develops into an offspring that inherits traits from each parent. By convention, organisms that produce smaller, more mobile gametes are called male, while organisms that produce produce larger, non-mobile gametes are called female. An organism that produces both types of gamete is hermaphrodite.

Sociobiology is a field of biology that aims to explain social behavior in terms of evolution. It draws from disciplines including psychology, ethology, anthropology, evolution, zoology, archaeology, and population genetics. Within the study of human societies, sociobiology is closely allied to evolutionary anthropology, human behavioral ecology, evolutionary psychology, and sociology.

<span class="mw-page-title-main">Evolution of sexual reproduction</span> How sexually reproducing multicellular organisms could have evolved from a common ancestor species

Sexual reproduction is an adaptive feature which is common to almost all multicellular organisms and various unicellular organisms. Currently, the adaptive advantage of sexual reproduction is widely regarded as a major unsolved problem in biology. As discussed below, one prominent theory is that sex evolved as an efficient mechanism for producing variation, and this had the advantage of enabling organisms to adapt to changing environments. Another prominent theory, also discussed below, is that a primary advantage of outcrossing sex is the masking of the expression of deleterious mutations. Additional theories concerning the adaptive advantage of sex are also discussed below. Sex does, however, come with a cost. In reproducing asexually, no time nor energy needs to be expended in choosing a mate and, if the environment has not changed, then there may be little reason for variation, as the organism may already be well-adapted. However, very few environments have not changed over the millions of years that reproduction has existed. Hence it is easy to imagine that being able to adapt to changing environment imparts a benefit. Sex also halves the amount of offspring a given population is able to produce. Sex, however, has evolved as the most prolific means of species branching into the tree of life. Diversification into the phylogenetic tree happens much more rapidly via sexual reproduction than it does by way of asexual reproduction.

In evolutionary biology, inclusive fitness is one of two metrics of evolutionary success as defined by W. D. Hamilton in 1964:

In biology, gonochorism is a sexual system where there are two sexes and each individual organism is either male or female. The term gonochorism is usually applied in animal species, the vast majority of which are gonochoric.

<span class="mw-page-title-main">Anisogamy</span> Sexual reproduction involving a large, female gamete and a small, male gamete

Anisogamy is a form of sexual reproduction that involves the union or fusion of two gametes that differ in size and/or form. The smaller gamete is male, a sperm cell, whereas the larger gamete is female, typically an egg cell. Anisogamy is predominant among multicellular organisms. In both plants and animals gamete size difference is the fundamental difference between females and males.

<span class="mw-page-title-main">Male</span> Sex of an organism which produces sperm

Male is the sex of an organism that produces the gamete known as sperm, which fuses with the larger female gamete, or ovum, in the process of fertilisation. A male organism cannot reproduce sexually without access to at least one ovum from a female, but some organisms can reproduce both sexually and asexually. Most male mammals, including male humans, have a Y chromosome, which codes for the production of larger amounts of testosterone to develop male reproductive organs.

In biology, altruism refers to behaviour by an individual that increases the fitness of another individual while decreasing their own. Altruism in this sense is different from the philosophical concept of altruism, in which an action would only be called "altruistic" if it was done with the conscious intention of helping another. In the behavioural sense, there is no such requirement. As such, it is not evaluated in moral terms—it is the consequences of an action for reproductive fitness that determine whether the action is considered altruistic, not the intentions, if any, with which the action is performed.

The evolution of human intelligence is closely tied to the evolution of the human brain and to the origin of language. The timeline of human evolution spans approximately seven million years, from the separation of the genus Pan until the emergence of behavioral modernity by 50,000 years ago. The first three million years of this timeline concern Sahelanthropus, the following two million concern Australopithecus and the final two million span the history of the genus Homo in the Paleolithic era.

Human behavioral ecology (HBE) or human evolutionary ecology applies the principles of evolutionary theory and optimization to the study of human behavioral and cultural diversity. HBE examines the adaptive design of traits, behaviors, and life histories of humans in an ecological context. One aim of modern human behavioral ecology is to determine how ecological and social factors influence and shape behavioral flexibility within and between human populations. Among other things, HBE attempts to explain variation in human behavior as adaptive solutions to the competing life-history demands of growth, development, reproduction, parental care, and mate acquisition. HBE overlaps with evolutionary psychology, human or cultural ecology, and decision theory. It is most prominent in disciplines such as anthropology and psychology where human evolution is considered relevant for a holistic understanding of human behavior.

Joan Roughgarden is an American ecologist and evolutionary biologist. She has engaged in theory and observation of coevolution and competition in Anolis lizards of the Caribbean, and recruitment limitation in the rocky intertidal zones of California and Oregon. She has more recently become known for her rejection of sexual selection, her theistic evolutionism, and her work on holobiont evolution.

The concept of the evolution of morality refers to the emergence of human moral behavior over the course of human evolution. Morality can be defined as a system of ideas about right and wrong conduct. In everyday life, morality is typically associated with human behavior rather than animal behavior. The emerging fields of evolutionary biology, and in particular evolutionary psychology, have argued that, despite the complexity of human social behaviors, the precursors of human morality can be traced to the behaviors of many other social animals. Sociobiological explanations of human behavior remain controversial. Social scientists have traditionally viewed morality as a construct, and thus as culturally relative, although others such as Sam Harris argue that there is an objective science of morality.

The evolutionary psychology of religion is the study of religious belief using evolutionary psychology principles. It is one approach to the psychology of religion. As with all other organs and organ functions, the brain's functional structure is argued to have a genetic basis, and is therefore subject to the effects of natural selection and evolution. Evolutionary psychologists seek to understand cognitive processes, religion in this case, by understanding the survival and reproductive functions they might serve.

<span class="mw-page-title-main">Hermaphrodite</span> Sexually reproducing organism that produces both male and female gametes

A hermaphrodite is a sexually reproducing organism that produces both male and female gametes. Animal species in which individuals are of different sexes, either male or female but not both, are gonochoric, which is the opposite of hermaphroditic.

<span class="mw-page-title-main">Female</span> Sex of an organism that produces ova

An organism's sex is female if it produces the ovum, the type of gamete that fuses with the male gamete during sexual reproduction.

Mary Jane West-Eberhard is an American theoretical biologist noted for arguing that phenotypic and developmental plasticity played a key role in shaping animal evolution and speciation. She is also an entomologist notable for her work on the behavior and evolution of social wasps.

The theoretical foundations of evolutionary psychology are the general and specific scientific theories that explain the ultimate origins of psychological traits in terms of evolution. These theories originated with Charles Darwin's work, including his speculations about the evolutionary origins of social instincts in humans. Modern evolutionary psychology, however, is possible only because of advances in evolutionary theory in the 20th century.

Inclusive fitness in humans is the application of inclusive fitness theory to human social behaviour, relationships and cooperation.

References

  1. 1 2 3 4 5 6 7 8 9 Roughgarden, Joan (2009). Genial Gene. California: The Regents of University of California. ISBN   978-0-520-25826-6.
  2. Roughgarden, Joan; Açkay, Erol. "Do we need a Sexual Selection 2.0?". The Association for the Study of Animal Behaviour.
  3. Roughgarden, Joan; Akçay, Erol (2009). "Final response:sexual selection needs an alternative". The Association for the Study of Animal Behaviour.
  4. 1 2 3 Atkinson, Nick (May 5, 2006). "Sexual selection alternative slammed". The Scientist . Archived from the original on 2007-09-29. Retrieved 2007-05-15.
  5. 1 2 3 Dall, S. R. X.; McNamara, J. M.; Wedell, N.; Hosken, D. J. (2006-05-05). "Debating Sexual Selection and Mating Strategies" (PDF). Science. 312 (5774): 689b–697b. doi:10.1126/science.312.5774.689b. PMID   16675684. S2CID   10887773 . Retrieved 14 June 2021.
  6. 1 2 West-Eberhard, Mary Jane (1975). "The Evolution of Social Behavior by Kin Selection". The Quarterly Review of Biology . 50 (1). University of Chicago Press: 1–33. doi:10.1086/408298. JSTOR   2821184. S2CID   14459515.
  7. 1 2 West-Eberhard, Mary Jane (1979). "Sexual Selection, Social Competition, and Evolution". Proceedings of the American Philosophical Society . 123 (4). American Philosophical Society: 222–34. JSTOR   986582.
  8. 1 2 West-Eberhard, Mary Jane (1983). "Sexual Selection, Social Competition, and Speciation". Quarterly Review of Biology . 58 (2). University of Chicago Press: 155–183. doi:10.1086/413215. JSTOR   2828804. S2CID   54711556.
  9. 1 2 West-Eberhard, Mary Jane (2014). "Darwin's forgotten idea: the social essence of sexual selection". Neuroscience & Biobehavioral Reviews . 46 (4). Elsevier: 501–508. doi:10.1016/j.neubiorev.2014.06.015. PMID   25003806. S2CID   1604935.
  10. Lyon, Bruce E.; Montgomerie, Robert (2012). "Sexual selection is a form of social selection". Philosophical Transactions of the Royal Society B . 367 (1600). London, UK: Royal Society: 2266–2273. doi:10.1098/rstb.2012.0012. PMC   3391428 . PMID   22777015.
  11. 1 2 Nesse, Randolph (2019). Good Reasons for Bad Feelings: Insights from the Frontier of Evolutionary Psychiatry. Dutton. pp. 172–176. ISBN   978-1101985663.
  12. Boehm, Christopher (1999). "The Natural Selection of Altruistic Traits" (PDF). Human Nature . 10 (3). Springer Science+Business Media: 205–252. doi:10.1007/s12110-999-1003-z. PMID   26196335. S2CID   207392341 . Retrieved July 7, 2021.
  13. Boehm, Christopher (2001) [1999]. Hierarchy in the Forest: The Evolution of Egalitarian Behavior (Revised ed.). Cambridge, MA: Harvard University Press. ISBN   978-0674006911.
  14. 1 2 Nesse, Randolph M. (2007). "Runaway social selection for displays of partner value and altruism". Biological Theory . 2 (2). Springer Science+Business Media: 143–55. doi:10.1162/biot.2007.2.2.143. S2CID   195097363.
  15. 1 2 Nesse, Randolph M. (2009). "10. Social Selection and the Origins of Culture". In Schaller, Mark; Heine, Steven J.; Norenzayan, Ara; Yamagishi, Toshio; Kameda, Tatsuya (eds.). Evolution, Culture, and the Human Mind. Philadelphia: Taylor & Francis. pp. 137–50. ISBN   978-0805859119.
  16. Boehm, Christopher (2012). Moral Origins: The Evolution of Virtue, Altruism, and Shame. New York: Basic Books. ISBN   978-0465020485.
  17. Boehm, Christopher (2014). "The moral consequences of social selection". Behaviour . 151 (2–3). Brill Publishers: 167–183. doi:10.1163/1568539X-00003143 . Retrieved July 7, 2021.
  18. Iyer, Priya; Roughgarden, Joan (2008). "Gametic Conflict Versus Contact in the Evolution of Anisogamy". Theoretical Population Biology. 73 (4): 461–72. doi:10.1016/j.tpb.2008.02.002. PMID   18485981.
  19. Iyer, Priya; Roughgarden, Joan (2008). "Dioecy as a Specialization Promoting Sperm Delivery". Evolutionary Ecology Research. 10.
  20. Roughgarden, Joan (2009). Evolution's Rainbow. University of California Press. pp. 129–131. ISBN   978-0520260122.
  21. Milam, Erika L.; Millstein, Roberta L.; Potochnik, Angela; Roughgarden, Joan E. (23 November 2010). "Sex and sensibility: The role of social selection". Metascience. 20 (2): 253–277. doi:10.1007/s11016-010-9464-6. S2CID   17700476.
  22. Clutton-Brock, Tim (2009). "We do not need a Sexual Selection 2.0-nor a theory of Genial Selection". The Association for the Study of Animal Behavior via Elsevier.
  23. Buss, David M. (1989). "Sex differences in human mate preferences: Evolutionary hypotheses tested in 37 cultures". Behavioral and Brain Sciences . 12 (1). Cambridge University Press: 1–49. doi: 10.1017/S0140525X00023992 .
  24. Buss, David M. (1995) [1992]. "5. Mate Preference Mechanisms: Consequences for Partner Choice and Intrasexual Competition". In Barkow, Jerome H.; Cosmides, Leda; Tooby, John (eds.). The Adapted Mind: Evolutionary Psychology and the Generation of Culture. New York: Oxford University Press. pp. 253–256. ISBN   978-0195101072.
  25. Miller, Geoffrey F. (2000). The Mating Mind: How Sexual Choice Shaped the Evolution of Human Nature (1st ed.). New York: Doubleday. pp. 292–340. ISBN   978-0385495165.
Bundled references

Further reading