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Unit of selection

In evolutionary biology, the unit of selection refers to the entity or level—such as genes, individual organisms, groups, or populations—that exhibits heritable variation, differential reproductive success, and the potential for evolutionary change through natural selection. This concept addresses how selection operates across biological hierarchies, with foundational criteria established by Richard Lewontin in 1970 requiring phenotypic variation among entities, differences in their fitness (survival and reproduction rates), and heritability of those fitness differences across generations. The debate over the unit of selection has evolved since Charles Darwin's era, when he primarily viewed individual organisms as the targets of selection, though he acknowledged potential group-level effects in social species like humans.^[1] In the mid-20th century, proponents of group selection, such as V. C. Wynne-Edwards, argued that natural selection could favor traits benefiting entire populations or groups, even at the expense of individual fitness, to explain phenomena like population regulation and altruism.^[2] However, this view faced strong criticism for lacking empirical support and mechanistic plausibility, as group-level adaptations would require rare conditions like frequent group extinctions and low within-group variation.^[1] A pivotal shift occurred in the 1960s and 1970s with the gene-centered perspective, advanced by George C. Williams and Richard Dawkins, who posited the gene as the fundamental unit of selection due to its long-term stability, capacity for replication across generations, and role in maximizing its own propagation via individual and extended phenotypes.^[1] Williams critiqued group selection as an inefficient and unnecessary explanation, emphasizing that adaptations evolve to enhance individual inclusive fitness rather than collective welfare, with group benefits often emerging as byproducts of individual actions.^[1] Dawkins further elaborated this in The Selfish Gene (1976), framing organisms as "vehicles" or survival machines built by genes to ensure their replication, thereby resolving apparent altruisms through kin selection and inclusive fitness concepts developed by W. D. Hamilton. Contemporary discussions recognize multilevel selection theory, integrating gene, individual, and group levels, where selection can act simultaneously across hierarchies but with varying efficacy depending on the context—such as in microbial communities or eusocial insects where group-level traits like colony defense confer fitness advantages.^[3] This framework, supported by empirical studies on phenomena like bacterial biofilms and human cooperation, underscores that no single level universally dominates, but the gene remains the ultimate beneficiary of selection due to its heritability.^[4] Ongoing research continues to refine these ideas, incorporating non-reproductive contexts like cultural evolution and ecological persistence to broaden the applicability of selection units beyond traditional biological lineages.^[4]

Overview

Definition

The unit of selection in evolutionary biology refers to the entity upon which natural selection acts directly, resulting in differential survival and reproduction that drives evolutionary change.^[5] This entity is the target of selection pressures, where variations among individuals lead to disparities in reproductive success, thereby altering the composition of future generations.^[2] A key distinction in understanding selection involves proximate and ultimate causation. Proximate causation addresses the immediate mechanisms—such as physiological or developmental processes—that enable an entity's response to its environment, explaining how selection occurs at the mechanistic level.^[6] In contrast, ultimate causation pertains to the evolutionary reasons why those mechanisms exist, rooted in the historical outcomes of natural selection that enhance fitness over generations.^[6] Within this framework, the distinction between replicators and interactors, introduced by Richard Dawkins and David Hull, highlights key aspects of selection. Replicators, such as genes, are the fundamental units that are faithfully copied and propagated across generations, serving as the beneficiaries of selection.^[5] Interactors, like organisms, act as cohesive wholes that interact with the environment, influencing the replication success of the genes they carry.^[5] This duality highlights how selection operates on observable entities while ultimately favoring the persistence of underlying genetic material. For an entity to qualify as a unit of selection, as established by Richard Lewontin (1970), it must demonstrate three essential properties: heritable variation, differential fitness, and the heritability of those fitness differences. Heritable variation ensures that traits passed to offspring can differ among entities, providing the raw material for selection.^[5] Differential fitness occurs when these variations lead to unequal reproductive success in response to environmental conditions.^[2] Finally, the heritability of fitness differences guarantees that advantageous traits are reliably transmitted, allowing selection to accumulate adaptive changes over time; this process, originating in Darwin's foundational work, is mathematically captured in models like the Price equation.^[7]^[8]

Historical Context

Charles Darwin's On the Origin of Species (1859) laid the foundational ideas of evolutionary theory by describing natural selection as a process acting primarily on individual organisms, where heritable variations in traits confer advantages in survival and reproduction, leading to the adaptation of populations over time. Although Darwin did not explicitly define a "unit of selection," his emphasis on inherited differences among organisms as the basis for evolutionary change implicitly positioned the individual as the key entity upon which selection operates.^[9] In the late 19th century, August Weismann advanced this framework with his germ-plasm theory, articulated in The Germ-Plasm: A Theory of Heredity (1893), which proposed a strict separation between the germline—carrying the hereditary material (germ-plasm)—and the somatic cells of the body.^[10] Weismann argued that only changes in the germ-plasm are heritable, effectively elevating genes or germinal elements as the true units of inheritance and selection, while somatic modifications acquired during an organism's lifetime cannot influence future generations.^[11] This theory resolved ambiguities in Darwin's ideas about inheritance and reinforced a shift toward viewing discrete hereditary units as central to evolution. The early 20th century saw the maturation of population genetics in the 1930s and 1940s, integrating Mendelian inheritance with Darwinian selection through the works of Ronald Fisher, J.B.S. Haldane, and Sewall Wright, which mathematically demonstrated how gene frequencies change under selection pressures. This synthesis shifted conceptual focus from organisms to genes as the fundamental units of selection, as models showed that adaptive evolution occurs via differential reproduction of alleles within populations.^[12] Within this emerging field, Haldane's The Causes of Evolution (1932) introduced a cost-benefit analysis of altruistic behaviors, calculating that such traits could evolve if the benefits to genetic relatives outweighed the costs to the actor, thereby foreshadowing later developments in understanding selection at familial levels. By the mid-20th century, debates intensified over the appropriate levels of selection, culminating in George C. Williams' Adaptation and Natural Selection (1966), which sharply critiqued group selection theories prevalent in earlier ecological interpretations.^[1] Williams argued that adaptations are best explained as outcomes of selection on individual genes or organisms, dismissing group-level benefits as incidental byproducts rather than causal drivers, thus solidifying the gene-centered view in evolutionary biology.^[13]

Theoretical Foundations

Core Principles

The unit of selection in evolutionary biology refers to the entity upon which natural selection acts, characterized by three essential conditions: variation in traits, heritability of those traits, and differential reproductive success. Heritability is a foundational requirement, meaning that for selection to occur, the variation among units must be reliably transmitted to subsequent generations with sufficient fidelity to allow cumulative change over time.^[14] This fidelity ensures that advantageous traits are not lost through random errors in replication, enabling the persistence and spread of beneficial variations. Without heritability, any differences in survival or reproduction would fail to produce evolutionary adaptation, as offspring would not inherit the features that conferred success to their parents.^[15] Differential reproduction forms the mechanism by which selection operates, where units exhibiting higher fitness—defined as their relative capacity to survive and reproduce—propagate more copies of themselves in the population compared to less fit variants.^[14] This process results in the increasing frequency of heritable traits that enhance fitness, driving evolutionary change at the level of the selected unit. Fitness here is not absolute but context-dependent, influenced by environmental interactions that favor certain variants over others.^[15] These principles apply universally across biological hierarchies, from genes to groups, where selection can act simultaneously at multiple levels provided the conditions of variation, heritability, and differential success are met at each.^[16] A key conceptual distinction clarifies the roles within this framework: replicators and interactors (or vehicles). Replicators are the entities whose information is copied with high fidelity during reproduction, such as genes, which serve as the fundamental units of inheritance. In contrast, interactors—or vehicles—are the higher-level structures, like organisms, that physically interact with the environment to affect the survival and replication of those replicators; organisms act as temporary carriers of genetic information, shielding and promoting its propagation without themselves being the primary targets of selection.^[17] This separation, first articulated by Dawkins in 1976 and refined by Hull in 1980, underscores that while interactors experience selection pressures directly, the long-term evolutionary legacy resides in the replicators they support.^[18] Maynard Smith further emphasized the vehicle concept in the 1980s as a disposable scaffold for replicator success, highlighting its role in models of evolutionary stability.

Mathematical Models

The Price equation provides a general mathematical framework for describing evolutionary change in the average value of a trait z across generations in a population, partitioning the total change \Delta \bar{z} into components due to selection and transmission. The full form of the equation is

\Delta \bar{z} = \frac{\text{Cov}(w, z)}{\bar{w}} + \frac{\text{E}(w \Delta z)}{\bar{w}},

where w is individual fitness (typically the number of offspring), \bar{w} is mean fitness, \text{Cov}(w, z) is the covariance between fitness and trait value capturing selection on the trait, and \text{E}(w \Delta z) is the expected change in the trait due to transmission biases weighted by fitness. This formulation, derived by George R. Price, holds as an exact identity under minimal assumptions about the mapping from parental to offspring generations, allowing analysis of how traits evolve through differential reproduction and inheritance fidelity.^[7] A key aspect of the Price equation's derivation involves its recursive structure, which enables partitioning of evolutionary change into selection within groups and between groups when populations are hierarchically structured. Starting from the basic covariance form, the equation can be expanded by considering group-level averages \bar{z}_g and within-group deviations z_i - \bar{z}_g, yielding a multilevel decomposition that quantifies how selection acts at different organizational levels without presupposing which level dominates.^[19] This partitioning reveals the conditions under which group-level traits can respond to selection, provided transmission maintains trait associations across levels. In social evolution, Hamilton's rule extends these ideas to identify conditions for the spread of altruistic traits, stating that a behavior evolves if rB > C, where r is the genetic relatedness between actor and recipient, B is the fitness benefit to the recipient, and C is the fitness cost to the actor. Derived from inclusive fitness considerations, this inequality applies the Price equation to kin-selected units by showing how relatedness amplifies benefits to shared genes, allowing identification of selectable social structures where direct fitness costs are offset. Hamilton's formulation thus formalizes how units beyond individuals, such as kin groups, can be vehicles for selection when genetic correlations align costs and benefits appropriately.^[20] The multilevel extension of the Price equation explicitly incorporates hierarchical selection, given by

\Delta \bar{z} = \frac{\text{Cov}(w_i, z_i)}{\bar{w}} + \frac{\text{Cov}(w_g, \bar{z}_g)}{\bar{w}},

where the first term represents within-group selection on individual deviations z_i, and the second term captures between-group selection on group averages \bar{z}_g with group fitness w_g. This form, building on Price's 1972 analysis, allows quantification of trade-offs between levels, such as when within-group competition undermines group-level adaptation. It demonstrates that multilevel selection occurs whenever variance in group traits correlates with group productivity, independent of single-level dynamics.^[19] Despite its generality, the Price equation relies on assumptions of trait additivity in multilevel partitions, where non-additive interactions (e.g., epistasis) can complicate unambiguous level attribution, and no transmission bias in the selection term, meaning the equation's first term isolates selection only if inheritance is faithful without systematic deviations.^[7] These constraints limit its direct application to complex systems with gene-environment interactions or frequent mutations altering heritability.^[21]

Levels of Selection

Molecular and Genetic Levels

At the molecular and genetic levels, the gene serves as the canonical unit of selection, where individual alleles compete for representation within genomes, often prioritizing their own replication over the organism's overall fitness. This perspective is encapsulated in the selfish gene hypothesis, which posits that genes act as the primary replicators driving evolutionary change through differential survival and transmission.^[22] In this framework, alleles within a genome engage in intragenomic competition, such as through mechanisms that bias inheritance to favor one variant over others.^[23] Selection operates directly on DNA sequences, the fundamental nucleic acid structures, through processes like mutation, recombination, and genetic drift, which collectively shape sequence variation and evolutionary trajectories. Mutations introduce novel variants, while recombination reshuffles existing sequences, and drift randomly alters allele frequencies, particularly in small populations; natural selection then favors sequences conferring advantages in replication or stability.^[24] A prominent example of intragenomic conflict at this level is meiotic drive, where certain alleles manipulate meiosis to achieve transmission rates exceeding the expected 50%, as observed in species like Drosophila and maize, leading to evolutionary arms races within the genome.^[25] Epigenetic modifications represent another layer of selectable units at the molecular level, involving heritable changes to gene expression without altering the underlying DNA sequence, such as DNA methylation, which adds methyl groups to cytosine bases to silence genes across cell divisions and generations. These modifications can evolve under selection when they enhance adaptive responses, like stress tolerance, by providing reversible regulatory mechanisms.^[26] A classic case is paramutation in plants, first discovered in maize (Zea mays) in the 1950s, where one allele induces a heritable epigenetic state in a homologous allele, altering pigmentation traits and demonstrating transgenerational inheritance of silenced states.^[27] In bacteria, horizontal gene transfer (HGT) enables entire gene clusters or operons to act as selectable units, allowing rapid acquisition of adaptive traits like antibiotic resistance or metabolic capabilities from distantly related organisms, thereby accelerating evolution beyond vertical inheritance.^[28] This process underscores how mobile genetic elements can propagate independently, subject to selection based on their conferred benefits in specific environments. Similarly, non-coding RNAs such as transfer RNA (tRNA) and ribosomal RNA (rRNA) have evolved under selection for structural and functional optimization; tRNA diversification, for instance, coevolved with aminoacyl-tRNA synthetases to expand the genetic code, while rRNA structures in the ribosome's peptidyl transferase center refined translation efficiency over billions of years.^[29]^[30] Intracellular selection further highlights conflict at the genetic level through selfish genetic elements, including transposons, which are DNA segments that replicate and insert copies within the genome, often at the host's expense by disrupting genes or increasing mutation rates. These elements spread via mechanisms like transposition during replication, evading suppression by host defenses, and can drive innovation by facilitating genomic rearrangements, though their proliferation is counterbalanced by selection for genomic stability.^[31] The Price equation provides a mathematical framework for partitioning such genetic changes into components attributable to selection and transmission biases at these molecular scales.^[32]

Cellular and Organismal Levels

In unicellular organisms such as bacteria and protists, the individual cell serves as the primary unit of selection, where heritable variations in traits like growth rate or metabolic efficiency directly influence reproductive success in changing environments.^[33] For instance, bacteria exhibit quorum sensing, a density-dependent communication system that coordinates behaviors such as biofilm formation or virulence factor expression, enabling collective responses that enhance survival without requiring multicellular structure.^[34] This emergent property illustrates how selection at the cellular level can favor cooperative signaling molecules, like autoinducers, that synchronize population-level adaptations to resource availability or threats.^[35] The transition to multicellularity, which emerged around 600 million years ago during the Ediacaran period, marked a shift where groups of cells evolved to function as integrated units, suppressing intra-group conflicts to prioritize organismal fitness.^[36] This evolutionary step allowed for specialization and larger body sizes, but it also introduced vulnerabilities, as seen in the selfish cell hypothesis, where uncontrolled cellular proliferation—manifesting as cancer—represents a reversion to unicellular-like autonomy that undermines the multicellular whole.^[37] Cancer arises when mechanisms enforcing cellular restraint fail, permitting "cheater" cells to exploit cooperative tissues for their own replication, a dynamic that highlights ongoing tension between cellular and organismal levels of selection.^[38] At the organismal level, natural selection acts on integrated phenotypic traits, such as morphology and physiology, that determine whole-organism viability and reproduction in specific ecological niches. A classic example is Charles Darwin's observations of Galápagos finches during his 1830s voyage, where beak shape variations adapted to seed size and availability drove differential survival during droughts, favoring deeper beaks for harder seeds and illustrating selection on organismal form as a cohesive unit.^[39] These adaptations, shaped by environmental pressures, underscore how organismal traits evolve through heritable changes that enhance overall fitness, rather than isolated cellular advantages. The soma-germ distinction, formalized by August Weismann in the late 19th century, reinforces organismal integrity by establishing a barrier that prevents somatic mutations—those arising in non-reproductive body cells—from being transmitted to offspring, ensuring that selection operates primarily on germline variations.^[11] This Weismann barrier isolates the hereditary material in germ cells, directing evolutionary change toward stable, organism-wide adaptations while limiting the long-term impact of somatic-level selfishness.^[40] Apoptosis, or programmed cell death, serves as a key mechanism to maintain organismal unity by selectively eliminating rogue cells that might otherwise disrupt multicellular cooperation, thereby countering potential cellular-level selection pressures. In evolutionary models of multicellularity, apoptosis evolved to enforce division of labor, such as in experimental systems where it promotes cluster stability by sacrificing peripheral cells for overall group propagation.^[41] This process ensures that selection favors organisms capable of suppressing intra-organismal conflicts, solidifying the organism as the dominant unit of inheritance and adaptation.^[42] At the social level, selection acts on behavioral traits that emerge from interactions among individuals, such as altruism and cooperation, where an individual's actions benefit others at a potential cost to itself.^[43] These traits can evolve if they enhance the fitness of social groups or kin networks, as seen in eusociality among insects, where non-reproductive workers support colony reproduction.^[43] In haplodiploid systems like those of bees and ants, females share higher relatedness with sisters (0.75) than with their own offspring (0.5), favoring worker sterility to promote sibling rearing over personal reproduction.^[43] This asymmetry, outlined in Hamilton's kin selection theory, explains the prevalence of female-biased eusociality in the Hymenoptera order.^[43] Group selection posits that traits benefiting the collective can spread if intergroup competition outweighs intragroup conflict, as proposed by Wynne-Edwards in his model of population regulation through social displays that limit breeding to prevent overexploitation of resources.^[44] Although initially criticized for lacking mechanistic detail, modern formulations revived the concept via trait-group models, where temporary assemblages of individuals allow between-group variance in trait composition to drive selection for altruism.^[45] In these models, altruistic behaviors persist if groups with more cooperators outcompete selfish ones, even as selfishness dominates within groups.^[45] Hamilton's rule, rB > C, briefly references how relatedness amplifies benefits in such contexts without requiring strict group benefits alone.^[43] Illustrative examples include alarm calls in vertebrates, such as those by Belding's ground squirrels, where callers risk predation to warn kin and group members, enhancing collective survival and potentially evolving via multilevel selection on both kin and group fitness components. In humans, tribalism—manifest as parochial altruism favoring ingroup cooperation and outgroup hostility—may represent a group-level adaptation shaped by intergroup competition in ancestral small-scale societies, promoting cultural norms that boost group productivity and defense.^[46] Within social systems, individual and group fitness often trade off: behaviors like worker foraging in eusocial colonies reduce personal reproduction but elevate colony output, with selection favoring those that maximize inclusive fitness across levels.^[47] Spatial structure, or population viscosity, further enhances group benefits by limiting dispersal, thereby increasing local relatedness and reducing the dilution of cooperative traits through mixing with selfish individuals from other groups.^[48] This viscosity creates "viscous" populations where interactions occur predominantly among relatives or familiar associates, amplifying the relative strength of group selection over individual-level pressures.^[49] Such dynamics underscore how social and group levels integrate organismal traits into emergent properties, resolving trade-offs through structured interactions that align individual sacrifices with collective gains.^[47]

Higher Taxonomic Levels

Species selection refers to the differential survival and proliferation of species based on their varying rates of speciation and extinction, acting as a higher-level analog to natural selection among individuals. This process, often termed "species sorting," posits that traits influencing a species' longevity, geographic range, or splitting rate can be selected if they enhance the production of daughter species or resistance to extinction. Steven M. Stanley formalized this concept in 1975, arguing that macroevolutionary patterns, such as trends in body size or diversity, arise not from gradual within-species change but from sorting among species with heritable differences in these demographic properties.^[50] At the clade level, selection can operate on entire lineages through major evolutionary transitions that create new units of selection, such as the origin of eukaryotic cells approximately 2 billion years ago. Eukaryogenesis, involving the endosymbiotic integration of mitochondria into an archaeal host, transformed simple prokaryotic life into complex cells capable of greater energy efficiency and multicellularity, representing a selectable innovation that propelled clade diversification.^[51] This transition, one of several identified by John Maynard Smith and Eörs Szathmáry, illustrates how clade-level selection favors emergent properties like compartmentalization that enhance heritability and adaptability across geological epochs. Punctuated equilibrium complements these ideas by emphasizing rapid speciation events as the primary units of macroevolutionary change, rather than slow, uniform transformation. Proposed by Niles Eldredge and Stephen Jay Gould in 1972, this model describes long periods of stasis within species punctuated by bursts of morphological innovation during cladogenesis, where new species form in peripheral isolates and spread, driving higher-level patterns observable in the fossil record.^[52] An illustrative example is the post-Cretaceous-Paleogene (K-Pg) diversification of birds, where avian clades underwent accelerated genomic and phenotypic evolution following the extinction of non-avian dinosaurs 66 million years ago, rapidly filling ecological niches with increased population sizes and brain-to-body ratios.^[53] Similarly, the Red Queen hypothesis, articulated by Leigh Van Valen in 1973, explains ongoing species-level co-evolution driven by biotic interactions, where lineages must continually adapt to counter coevolving antagonists, maintaining diversity through perpetual arms races at taxonomic scales. Despite these mechanisms, selection at higher taxonomic levels faces significant limitations, particularly weak heritability due to the vast geological timescales involved, which obscure direct transmission of species traits across generations. Traits like speciation rate may persist in daughter lineages but are diluted by stochastic events such as mass extinctions or environmental shifts, reducing the predictability and observability of macroevolutionary responses compared to lower levels.^[54]

Debates and Modern Developments

Gene-Centered Perspectives

The gene-centered perspective in evolutionary biology posits that genes, rather than organisms or groups, are the fundamental units of selection because they are the stable, heritable replicators that persist across generations. In this view, organisms function as transient vehicles that facilitate gene replication but are ultimately disposable, as their survival and reproduction serve the propagation of genetic material. This framework, articulated by Richard Dawkins, emphasizes that natural selection acts primarily on differential gene replication success, with phenotypic traits emerging as effects of genes rather than ends in themselves.^[17]^[55] A key pillar of this perspective is the concept of inclusive fitness, introduced by W. D. Hamilton, which reframes apparent altruism or group benefits as outcomes of gene-level accounting. Inclusive fitness measures the impact of a gene on its own transmission, including effects on relatives weighted by genetic relatedness, allowing selfish genes to explain cooperative behaviors without invoking higher-level selection. For instance, Hamilton's rule (rb > c) demonstrates how genes promoting costly aid to kin can spread if the inclusive fitness benefit exceeds the direct cost, resolving puzzles like eusociality in insects through gene frequency changes rather than organismal or group optimality. This gene-focused lens integrates individual and kin selection into a unified theory of genic propagation.^[56] Empirical support for gene primacy comes from genome-wide association studies (GWAS) that reveal how specific alleles influence fitness components, such as survival and reproduction, driving evolutionary change through their frequency shifts. For example, analyses of human genetic variation have estimated allele-specific fitness effects across protein-coding regions, showing that variants with positive selection signatures correlate with adaptive traits at the genic level. Similarly, selfish genetic elements, like B chromosomes in various species, provide direct evidence of intragenomic conflict where non-essential DNA sequences bias their own transmission at the expense of host fitness, underscoring selection among genes within the genome. These elements, observed in organisms from plants to mammals, accumulate despite reducing organismal viability, confirming that genes can act as independent replicators.^[57]^[58]^[31]^[23] Apparent organismal adaptations, such as complex morphological traits, are thus interpreted as byproducts of cumulative gene frequency changes under selection, rather than direct optimizations at the individual level. Post-2000 genomic advancements, including whole-genome sequencing, have bolstered this view by documenting intragenomic selection dynamics, such as meiotic drive in selfish elements and positive selection on specific loci, revealing how gene-level competition shapes evolutionary outcomes without requiring multilevel explanations. This historical emphasis on genes traces back briefly to August Weismann's germ plasm theory, which isolated hereditary material in germ cells, laying groundwork for viewing genes as the enduring focus of inheritance.^[31]^[11]

Multilevel Selection Advocacy

Advocates of multilevel selection (MLS) emphasize that natural selection can operate effectively at multiple hierarchical levels simultaneously, rather than being confined to a single unit such as the gene or individual. A key conceptual distinction in this advocacy is between MLS1 and MLS2, as formulated by Damuth and Heisler. In MLS1, group-level change arises from contextual differences in individual fitness across groups, where group fitness is simply the average of individual fitnesses within that context. In contrast, MLS2 treats groups as having emergent properties that confer independent fitness to the group as a whole, allowing for genuine group-level adaptation independent of individual-level effects.^[59] This framework enables the partitioning of evolutionary change across levels, highlighting how selection pressures can align or conflict hierarchically. Sober and Wilson advanced MLS advocacy through their trait-group model, detailed in their 1998 book Unto Others: The Evolution and Psychology of Unselfish Behavior. They argued that group-level adaptations, such as altruism and moral sentiments that benefit the collective, can evolve when between-group variance in fitness exceeds within-group variance, even if individual-level selection favors selfishness. This model reconciles individual and group benefits by viewing groups as temporary assemblages where traits like cooperation enhance group survival and reproduction, thereby indirectly benefiting carriers of those traits. Their work provides a rigorous defense against criticisms of group selection, positing that unselfish behaviors in humans and other species are plausibly outcomes of multilevel processes.^[60] Empirical support for MLS comes from experimental evolution studies, particularly in microbial systems. For instance, Ratcliff et al. demonstrated between-group selection in yeast by selecting for faster settling in liquid media, leading to the rapid evolution of multicellular "snowflake" clusters within 60 generations. These clusters exhibited division of labor and size-based fitness advantages at the group level, where larger aggregates outcompeted smaller ones and suppressed individual cheater cells that would otherwise disrupt cohesion, illustrating how group-level selection can drive the emergence of complexity.^[61] The major transitions framework further bolsters MLS by explaining how new levels of selection arise from lower ones. Maynard Smith and Szathmáry outlined eight key transitions in evolution, such as the shift from prokaryotic cells to eukaryotic ones and from unicellular to multicellular organisms, where cooperation at higher levels evolves by resolving conflicts at lower levels through mechanisms like genetic relatedness or policing. In the transition to multicellularity, for example, selection favors cell groups that maintain integrity over proliferating individuals, establishing the multicellular entity as a new unit of selection. In the 2010s, MLS advocacy extended to human evolution through models of parochial altruism, where individuals cooperate preferentially within their group while showing aggression toward out-groups. García and Bergh showed via simulations of a prisoner's dilemma with intergroup conflict that parochial strategies—altruistic in-group behavior combined with out-group hostility—can stably evolve under multilevel selection, as group success in competition amplifies the spread of such traits despite within-group costs. This mechanism has been proposed to explain the evolution of human cooperation in tribal contexts.^[62] The Price equation, by partitioning covariance between traits and fitness across levels, underscores how such multilevel dynamics contribute to overall evolutionary change. Gene-centered perspectives, while emphasizing selfish genes, can be integrated as a subset of MLS when viewed through hierarchical lenses.

Extensions to Cultural Evolution

The concept of the unit of selection extends beyond biological entities to cultural phenomena through memetics, where Richard Dawkins introduced the term "meme" in 1976 as a basic unit of cultural transmission analogous to genes, capable of replication, variation, and selection via imitation.^[22] Memes, such as ideas, behaviors, or styles, propagate through human minds and media, undergoing evolutionary processes that favor those conferring advantages in cultural fitness, like widespread adoption or memorability.^[22] Susan Blackmore expanded this framework in 1999, arguing that memes drive human evolution by selecting for imitation abilities, positioning memetics as a mechanism for explaining complex cultural traits like language and technology without relying solely on genetic inheritance.^[63] Dual inheritance theory further integrates cultural units into evolutionary models, positing that genes and culture co-evolve as parallel inheritance systems, with cultural traits transmitted vertically (from parents), horizontally (between peers), or obliquely (from non-parents).^[64] Developed by Robert Boyd and Peter Richerson in 1985, this theory uses mathematical models of cultural transmission—such as biased transmission where individuals preferentially adopt advantageous cultural variants—to demonstrate how culture can evolve independently yet interact with genetic selection, accelerating adaptation in variable environments.^[64] For instance, cultural practices like tool use can spread rapidly across populations, influencing genetic fitness by altering selective pressures. Specific examples illustrate cultural units as selectable entities; the evolution of language serves as a prime case, where linguistic structures—such as grammatical rules or vocabulary—undergo selection through cultural transmission, favoring learnable and communicative forms that persist across generations.^[65] Similarly, the spread of agriculture around 10,000 BCE exemplifies a selectable cultural innovation, as farming techniques propagated via imitation and migration, enabling population growth and reshaping human societies despite initial costs like labor intensity.^[66] Gene-culture coevolution highlights reciprocal selection between cultural practices and genetic traits, notably in lactase persistence, a genetic adaptation allowing adult milk digestion that arose after the introduction of dairy farming around 7000 BCE in Europe and Africa.^[67] This mutation spread rapidly in pastoralist populations because the cultural practice of dairying created a nutritional niche favoring the allele, demonstrating how cultural innovations can drive genetic change and vice versa.^[67] Digital memes, which first emerged in the late 1990s, have become increasingly potent cultural units in the 2020s, evolving through online platforms where they replicate via sharing, mutate through remixing, and face selection based on virality metrics like engagement rates.^[68] AI-generated content, including memes and viral ideas, represents a novel extension as of 2025, accelerating cultural evolution by automating variation and dissemination—such as through tools like DALL-E and Midjourney—though critiques emphasize lower fidelity in replication compared to biological genes, as digital memes often degrade or hybridize rapidly during transmission.^[69]^[70]^[71] Recent studies as of 2025 highlight how AI-assisted meme creation is shaping online communication and cultural trends, further blurring lines between human and machine-driven evolution.^[72]

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In this paper, I will argue that the generality of the Price equation comes at a cost, and that is that the terms in it become meaningless.
[21]
The Selfish Gene - Richard Dawkins - Oxford University Press
The Selfish Gene has become a classic exposition of evolutionary thought. Professor Dawkins articulates a gene's eye view of evolution.Missing: source | Show results with:source
[22]
Selfish genetic elements - PMC
George Williams' Adaptation and Natural Selection (1966) and Richard Dawkins' The Selfish Gene (1976) were instrumental in introducing the gene's-eye view to ...
[23]
DNA sequence diversity and the efficiency of natural selection in ...
Nov 9, 2016 · We explore and quantify the relationship between neutral genetic diversity (synonymous site diversity (π S )) and the efficiency of selection.
[24]
Meiotic drive mechanisms: lessons from Drosophila - Journals
Oct 23, 2019 · The resulting intragenomic conflicts triggered by meiotic drive create evolutionary arms races and shape genome evolution. The phenomenon of ...Abstract · Introduction · Consequences of drive · Conclusion: emerging themes...
[25]
Epigenetics and gene expression | Heredity - Nature
May 12, 2010 · Epigenetic processes, including DNA methylation, histone modification and various RNA-mediated processes, are thought to influence gene expression.
[26]
Paramutation phenomena in plants - PubMed
Aug 31, 2015 · Paramutation is a particular epigenetic phenomenon discovered in Zea mays by Alexander Brink in the 1950s, and then also found in other plants and animals.
[27]
Horizontal Gene Transfer and the Evolution of Bacterial and ... - NIH
Jan 15, 2013 · Here we synthesize recent literature demonstrating the importance of habitat and niche in structuring horizontal gene transfer.
[28]
Evolution of transfer RNA and the origin of the translation system - NIH
The evolutionary pattern of diversification of tRNA may indicate how the chemical relationship between these molecules and aminoacyl-tRNA synthesases could have ...
[29]
Tracing the evolution of RNA structure in ribosomes - Oxford Academic
The evolution of the complete repertoire of structural ribosomal characters was formally traced lineage-by-lineage in the tree, showing a tendency towards ...
[30]
Selfish genetic elements, genetic conflict, and evolutionary innovation
Jun 20, 2011 · An example will illustrate the point. A meiotic drive allele reduces the nondriving allele among the gametes of heterozygous individuals; ...Selfish Genetic Elements... · Types Of Genetic Conflict · Types Of Sges And Their...<|separator|>
[31]
Mutation and the evolution of recombination - PMC - NIH
Random drift is a more general source of negative linkage disequilibria, and can cause selection for recombination even in large populations, through the chance ...
[32]
On The Evolution of Bacterial Multicellularity - PMC - NIH
Jan 16, 2015 · Here we discuss examples of multicellular behaviors in bacteria, the selective pressures that may have led to their evolution, possible origins and ...
[33]
Workshop Overview | The Social Biology of Microbial Communities
According to Greenberg, quorum sensing and response is generally used to describe a cell-to-cell communication system that allows bacteria to monitor population ...
[34]
Cell Communications among Microorganisms, Plants, and Animals
The membrane receptors in bacteria often directly respond to nutrients [1], or to quorum sensing (QS) signaling molecules [2], or to various other molecules in ...
[35]
How Did Multicellular Life Evolve? | News | Astrobiology
Feb 13, 2017 · More complex forms of life took longer to evolve, with the first multicellular animals not appearing until about 600 million years ago. The ...
[36]
Cooperation and conflict in the evolution of multicellularity - Nature
Jan 1, 2001 · The affect of mutation on group size depends on whether mutant cells replicate faster (selfish) or slower (uniformly deleterious) than non- ...
[37]
[PDF] Cancer tumors as Metazoa 1.0: tapping genes of ancient ancestors
Feb 7, 2011 · By contrast, multicellularity demands that individual cells subordinate their selfish agenda to the requirements of the organism as a whole. In ...
[38]
Community-wide genome sequencing reveals 30 years of Darwin's ...
Sep 29, 2023 · This study takes advantage of 30 years of study of a classic system to elucidate the role of genetic architecture and introgression in adaptation.
[39]
Inheritance of somatic mutations by animal offspring - Science
Aug 31, 2022 · In 1892, Weismann first proposed the theory that somatic mutations acquired during an animal's lifetime are evolutionarily irrelevant because ...Missing: distinction | Show results with:distinction
[40]
Experimental evolution of multicellularity - PMC - PubMed Central
Jan 17, 2012 · We observed the evolution of division of labor through programmed cell death (apoptosis), a mechanism used by snowflake yeast to increase ...
[41]
Emergence of multicellularity in a model of cell growth, death and ...
Abstract. How multicellular life forms evolved from unicellular ones constitutes a major problem in our understanding of the evolution of our biosphere.
[42]
The genetical evolution of social behaviour. I - ScienceDirect.com
July 1964, Pages 1-16. Journal of Theoretical Biology. The genetical evolution of social behaviour. I. Author links open overlay panel W.D. Hamilton. Show more.
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Animal Dispersion in relation to social behaviour - Internet Archive
Mar 1, 2023 · Animal Dispersion in relation to social behaviour. by: V.C. Wynne-Edwards. Publication date: 1962. Publisher: Oliver and Boyd.
[44]
A theory of group selection. - PNAS
A simple model shows that this can lead to the selection of altruistic traits that favor the fitness of the group over that of the individual.
[45]
[PDF] Tribal Social Instincts and the Cultural Evolution of Institutions to ...
We argue that cultural group selection and related cultural evolutionary processes had an important role in shaping the innate components of our social ...
[46]
The group covariance effect and fitness trade-offs during ... - PNAS
This paper studies the role of fitness trade-offs in fitness reorganization, the evolution of cooperation, and the conversion of a group into a new individual ...
[47]
[PDF] Population Viscosity and the Evolution of Altruism
Under these conditions, population viscosity can support the selection of both weak and strong altruism.
[48]
[PDF] Genetic relatedness in viscous populations
Selection on social interactions depends on both the fitness consequences of the behaviour and on the relatedness of the actor to the individuals affected.
[49]
A theory of evolution above the species level. - PNAS
Feb 15, 1975 · Gradual evolutionary change by natural selection operates so slowly within established species that it cannot account for the major features of evolution.Missing: URL | Show results with:URL
[50]
The emerging view on the origin and early evolution of eukaryotic cells
Sep 11, 2024 · In a process referred to as eukaryogenesis, the eukaryotic cell is believed to have evolved between approximately 1.8 and 2.7 billion years ago ...Missing: date source
[51]
(PDF) Punctuated Equilibria: An Alternative to Phyletic Gradualism
In their 1972 paper, Eldredge and Gould devoted considerable space to a ... We based our "correct" ideas about punctuated equilibrium on the concepts that Gould ...
[52]
Genome and life-history evolution link bird diversification to the end ...
Jul 31, 2024 · We show that temporal proximity to the K–Pg boundary increases the probability of molecular model shifts (Fig. 1 and fig. S1, A to D), linking a ...
[53]
Heritability at the ecosystem level - PNAS
In a sexually reproducing species, individual selection can act only on the average effects of individual genes. ... limits the heritability of group- and ...
[54]
Selfish genetic elements and the gene's-eye view of evolution - PMC
The distinction between replicator and vehicles (Dawkins 1982a, 1982b; also known as interactors, Hull 1980) was introduced partly to address this issue.
[55]
Inclusive fitness is an indispensable approximation for ...
Hamilton (1964) showed that, at equilibrium, organisms should appear to be choosing traits with regards to inclusive fitness, and that this results from gene ...
[56]
Estimation of allele-specific fitness effects across human protein ...
Abstract. A central challenge in human genomics is to understand the cellular, evolutionary, and clinical significance of genetic variants.
[57]
Analyses of allele age and fitness impact reveal human beneficial ...
Jan 29, 2024 · Analyses of allele age and evolutionary impact reveal that beneficial alleles in a human population are often older than neutral controls.<|separator|>
[58]
https://elifesciences.org/reviewed-preprints/93258
[59]
Unto Others - Harvard University Press
Oct 1, 1999 · In Unto Others philosopher Elliott Sober and biologist David Sloan Wilson demonstrate once and for all that unselfish behavior is in fact an important feature.
[60]
https://www.hup.harvard.edu/books/9780674930476
[61]
https://doi.org/10.1073/pnas.1115323109
[62]
The Meme Machine - Susan Blackmore - Oxford University Press
The Meme Machine shows that once our distant ancestors acquired the crucial ability to imitate, a second kind of natural selection began.
[63]
Culture and the Evolutionary Process, Boyd, Richerson
Boyd and Richerson explore how genetic and cultural factors interact, under the influence of evolutionary forces, to produce the diversity we see in human ...
[64]
Language as an evolutionary system - ScienceDirect.com
Language is an evolutionary system because its transmission is the basis for its evolution, and it adapts to aid its own survival.
[65]
The Origins of Agriculture as a Natural Experiment in Cultural ...
Vavilov and Sauer argued that agriculture originated in locations where the wild ancestors of later crop species attracted the attention of hunters and ...
[66]
Evolution of lactase persistence: an example of human niche ...
This supports the idea that LP coevolved with the cultural adaptation of dairying as a gene–culture coevolution process. Nonetheless, the correlation between LP ...
[67]
Memes in a Digital World: Reconciling with a Conceptual ...
Apr 1, 2013 · The term meme was introduced by the biologist Richard Dawkins in his book The Selfish Gene (1976). As part of his larger effort to apply ...<|control11|><|separator|>
[68]
Entropy and complexity unveil the landscape of memes evolution
Oct 8, 2021 · Our study starts from Dawkins' hypothesis of the meme as the basic unit of cultural evolution, in connection with the post-memetics analyses of ...Results · Methods · Clustering
[69]
The Case for Memes | Biological Theory
Mar 20, 2015 · In this paper I identify the key objections to memetics as a viable explanatory tool in studies of cultural evolution. I attempt to defuse these ...Missing: critiques | Show results with:critiques