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Richard Dawkins: Genes, Phenotypes, and Memes

From The Selfish Gene to The Extended Phenotype and The Meme Theory: how evolution, behavior, and culture can be reinterpreted through…

Mattia Paparo in Sharing Science · 2026-06-26 19:20 · 169 claps · 11.7 min read paywalled
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Richard Dawkins: Genes, Phenotypes, and Memes

From The Selfish Gene to The Extended Phenotype and The Meme Theory: how evolution, behavior, and culture can be reinterpreted through replicators

Matthias Asgeirsson (2006). Richard Dawkins giving a lecture based on The God Delusion, Reykjavik. Source: Flickr / Wikimedia Commons. Licensed under CC BY-SA 2.0.

Matthias Asgeirsson (2006). Richard Dawkins giving a lecture based on The God Delusion, Reykjavik. Source: Flickr / Wikimedia Commons. Licensed under CC BY-SA 2.0.

Anyone who has studied biology or has in some way approached topics such as biological evolution cannot fail to know Richard Dawkins. He has been considered the “nemesis” of Stephen J. Gould, another outstanding evolutionary biologist, geologist, and paleontologist who, during the twentieth century, revolutionized — together with Dawkins himself — the way biological evolution is understood, interpreted, and studied. He is also known in the United States for his positions against religion in general, or against religious groups, and unfortunately also because his name reportedly appeared in the well-known Epstein Files.

I will not address that issue here; in this discussion, we will focus solely on his work and on several concepts of major importance in evolutionary biology (**here, if desired, further information regarding the matter and his alleged involvement can be found). In particular, I will discuss three topics that are highly relevant to the study of Human Ecology, Niche Theory**, and biological evolution in general.

Zoe Margolis (2009), Ariane Sherine and Richard Dawkins at the Atheist Bus Campaign launch, CC BY 2.0, Wikimedia Commons.

Zoe Margolis (2009), Ariane Sherine and Richard Dawkins at the Atheist Bus Campaign launch, CC BY 2.0, Wikimedia Commons.

Who is the evolutionary biologist Richard Dawkins?

Let us take a look at him as a person before delving into his books. He is a distinctive figure, just as distinctive are his studies.He was born in Nairobi on March 26, 1941, into an English family, and later moved to the United Kingdom. He studied at the University of Oxford, where he graduated in biology, and later pursued his PhD under the supervision of the Dutch ethologist Nikolaas Tinbergen. Subsequently, he moved to the United States, first to the University of Berkeley, before returning to Oxford, where he became a university professor teaching zoology. Since becoming a professor, he has written numerous essays and books focusing on genes, as he adopts a “gene-centered” view of life. This perspective can already be appreciated in The Selfish Gene, his first published work.

This line of thought perfectly embodies and reflects the concept of evolution, where change (evolution) occurs through evolutionary mechanisms such as natural selection, sexual selection, and others. Dawkins applies the concept of evolution directly to genes. If animals, plants, and organisms undergo genetic changes over time through random processes, then it is likely that genes themselves are also subject to selective and evolutionary mechanisms, since they possess a particular characteristic: they replicate themselves (replicators), doing so in order to “survive” across successive generations and for as long as possible.

Nicosmos (2011). Photograph of the French edition of The Selfish Gene (chapter “Memes: the new replicators”) by Richard Dawkins. Source: Wikimedia Commons. Licensed under CC BY-SA.

Nicosmos (2011). Photograph of the French edition of The Selfish Gene (chapter “Memes: the new replicators”) by Richard Dawkins. Source: Wikimedia Commons. Licensed under CC BY-SA.

At the center of evolutionary change lies the gene, and this is well established since genes mutate and are selected through various mechanisms in order to be transmitted to offspring. With a bit of luck, they may become more frequent within a population, thereby genetically altering it. However, Dawkins also sees something more: he views human beings and all organisms as “vehicles” that transport genes. Richard describes these vehicles in The Selfish Gene as a biological structures that evolved to ensure the survival and transmission of genes, which represent the fundamental units of genetic inheritance. At this point, the behaviors observed in social animals come into play, placing the discussion within both an ethical and ethological context. Behavior is encoded by genes, and the gene itself — rather than the individual organism — is identified as the primary unit of natural selection driving the evolutionary process.

According to this evolutionary perspective, the primary level at which natural selection acts is not the species, the group, or the individual, but rather the gene — considered the fundamental element of hereditary transmission. This leads to the assumption that the most important aspect of evolution is the good of the species (or the group), rather than the good of the individual (or the gene). This helps explain “unusual” behaviors such as the social organization of ants, or all those behaviors that contribute more to the well-being of the population than to the well-being of the individual itself. These are altruistic behaviors driven by another underlying behavior: a selfish one, namely the transmission of genes to future generations, to the next “vehicles” existing within a population of a species rather than solely within a single organism.

Illustration of social organization of ants. Photo by Marco Neri on Unsplash

Illustration of social organization of ants. Photo by Marco Neri on Unsplash

Naturally, one can imagine the many scientific debates and disagreements he encountered in the academic world, including those with his colleague, the paleontologist and evolutionary biologist Stephen J. Gould. Indeed, these “long-distance disputes” indirectly caused many people to misunderstand and distance themselves from the complex world of evolution and the emerging framework of modern evolutionary theory. This remains an issue affecting a substantial proportion of Americans who “do not believe in evolution”.

Dawkins later stated, after Gould’s death, that they could easily have collaborated, since their studies and ideas were not fundamentally in conflict (it should be remembered that Gould was one of the founding figures of modern evolutionary theory, having studied evolutionary mechanisms such as exaptation and punctuated equilibrium). Such a collaboration, according to Dawkins, could have benefited modern evolutionary theory considerably. A brief note: by “modern evolution”, I am referring broadly to all post-Darwinian studies without going into detail, otherwise it could create confusion.

Eric Taylor (1985). Book cover of Stephen Jay Gould, The Flamingo’s Smile. Source: Wikimedia Commons. Public Domain.

Eric Taylor (1985). Book cover of Stephen Jay Gould, The Flamingo’s Smile. Source: Wikimedia Commons. Public Domain.

His studies and theories also attempt to explain how life itself may have originated. In fact, he considers the RNA molecule to be the first “replicator.” The earliest biological components of the cell may have initially performed a protective function for replicators within this form of natural selection operating at the genetic level, eventually leading to the formation of the first true cells and, consequently, increasingly complex vehicles.

Dawkins is a committed atheist with a strong personality, and consequently his major critics have often been anti-evolutionists and religious groups. This is because his studies have indirectly challenged religious perspectives, as can be seen in his book The God Delusion, where he argues that religious beliefs may have a biological and evolutionary explanation. This idea is explored further in his article Viruses of the Mind, where he suggests that meme theory (according to which culture, similarly to evolutionary models, changes over time and is “transmitted” across generations through so-called memes) may provide an evolutionary explanation for religions.

Let us now go into detail by discussing the two most important essays and the theory of memes. These will be essential for a better understanding of niche theory and all topics related to Human Ecology.

Illustration of an mRNA molecule. Powered by Canva

Illustration of an mRNA molecule. Powered by Canva

THE SELFISH GENE (1976)

We are survival machines — self-propelled robots blindly programmed to preserve those selfish molecules known as genes

(Richard Dawkins, The Selfish Gene, Preface, 1976; translation based on the Italian edition)

His gene-centered view of life will be the core of the book, as every chapter depends on this concept. In this work, Dawkins identifies the gene, rather than the individual (or the species), as the unit of natural selection. This book has often been criticized due to misunderstandings, since genes are entities that replicate, but this does not imply that they are driven by motives or intentions; rather, it simply means that their effects can be accurately described as if they were. This is a simplification intended to make the concept easier to understand. The genes that are transmitted are only those whose consequences serve their own “interests”, extending beyond the individual or the species, and reaching the level of the population. Genes replicate in order to continue “surviving” across generations, seeking the best strategy to maximize their chances of replication.

With this study, many animal behaviors are analyzed using game theory, the best explanation assuming that the advantage lies with individual genes rather than the traditional interpretation centered on the benefit of individuals and species. This explains the concept of individual “altruism” in nature, especially in kin relationships, when an individual sacrifices its own life to save that of its relatives. The individual is “acting” in the interests of its genes. The universe is populated by stable things, such as a set of atoms, and from this consideration Dawkins develops the concept of a “replicator”: a stable molecule with the ability to replicate itself. This may have occurred in the primordial soup, a broth rich in replicators such as RNA (the probable first replicator). Replication is not always faithful, and sometimes incorrect copies arise, and it is thanks to these errors that changes (evolution) occurred, leading to the emergence of biodiversity.

Cover illustration of *The Selfish Gene* (Italian edition) called “Il Gene Egoista”. Personal photo.

Cover illustration of The Selfish Gene (Italian edition) called “Il Gene Egoista”. Personal photo.

Competition among these replicators and environmental pressures randomly selected the most suitable replicators, refining their propagation strategies. Some of them, according to Dawkins, may have evolved in such a way as to be able to “feed” on other replicators, while others may have developed a form of protective shield for defense, giving rise to the earliest primordial cells. The replicators of today are the genes contained within living organisms (which Dawkins calls “vehicles”). Genetic combinations, and therefore cooperation, increase the chances of survival, replication, and dissemination.

Organisms behave altruistically against their own individual interests (in terms of health, safety, and personal reproduction) in order to help related organisms reproduce. Thus, parental behaviors can be explained as if the gene were helping copies of itself replicate in other bodies. In this way, gene selfishness is translated into kin altruism among organisms. The important point is to ensure genetic continuity, and this can lead to the sacrifice of an individual for the good of the population. Dawkins’ perspective does not exclude modern evolutionary theory at all, as it only changes the viewpoint, offering a different way of seeing evolution. By reading the book, one understands that this perspective helps to further clarify evolutionary processes.

Illustration of an orangutan with a small individual it is trying to protect.. Photo by Joshua J. Cotten on Unsplash

Illustration of an orangutan with a small individual it is trying to protect.. Photo by Joshua J. Cotten on Unsplash

THE EXTENDED PHENOTYPE (1982)

If we wish, we can consider this book as a sequel to The Selfish Gene, since in the first we focus largely on the gene itself (the unit of heredity), while in this one we focus on the concept of phenotype (that is, how a gene is externally expressed). We must always remember that Dawkins’ gene-centered view of life is also present in this book; therefore, it is not a book that simply describes what a phenotype is, but rather how these selfish/altruistic genes are expressed, thereby completing his earlier work.

The essay presents itself as a development of the theory previously introduced in 1976, and much of the text serves as a discussion of the subsequent criticisms of that theory, namely evolution seen from the perspective of the gene, the entity on which natural selection would act (rather than on the organism), with organisms functioning only as “vehicles,” survival machines for genes, providing them with a means to be transmitted and to persist across generations.

Illustration of my personal copy of “The Extended Phenotype” in italian

Illustration of my personal copy of “The Extended Phenotype” in italian

In this book we see even more clearly the “downgrading” of the organism as a genetic vehicle, since an animal has the ability to influence its environment and other animals to its own advantage. The phenotype is the external expression of the gene, and we should not expect only a morphological coding (appearance) or only a behavioral one (why an animal behaves in a certain way), but rather we must adopt a deeper level of observation, which Dawkins illustrates with the following example: beavers possess genes that code for teeth (first phenotype), and thanks to these they can eat and defend themselves (second phenotype).

The extended phenotype is an extension, a way of looking beyond these two “classical” phenotypes, since beavers, through those teeth, build dams that lead to the formation of a lake. We can therefore consider the lake and the dam as “extended phenotypes”, elements determined by the same genes that code for teeth, and elements that can provide advantages by increasing the probability of the gene being replicated in the next generation.

Illustration of a beaver near a dam built close to a lake.. Photo by Howard Walsh on Unsplash

Illustration of a beaver near a dam built close to a lake.. Photo by Howard Walsh on Unsplash

One might wonder whether lakes and dams are indeed indirect consequences, but Dawkins explains that even beaver teeth are indirect in the same sense, since a single gene does nothing more than synthesize a protein, and from that protein the influence on tooth structure emerges through intermediate steps. These steps are numerous and extremely complex, involving the action of many genes. In fact, it is extremely rare, if not almost impossible, to find a single gene coding for a single trait; it is more common to find multiple genes that “cooperate” and influence one another.

The organism is always and only considered a genetic vehicle, an external realization of genes used by them to compete with one another. “Extended” consequences can be such that they induce an organism that does not possess those specific genes to exploit those same extensions. We are fully in the field of parasitism, and Dawkins cites the specific example of the ant Monomorium santschii, whose queen infiltrates colonies of other species, inducing (probably chemically) the worker ants to kill their own queen and take her place. Monomorium benefits from “extended phenotypes” coded by genes it does not possess, while this behavior is harmful to the genes of those ants, because the only fertile individual capable of perpetuating those shared genes is the queen. Dawkins summarizes these findings in what he calls the Central Theorem of the Extended Phenotype, stating that the behavior of an animal tends to maximize the survival of the genes responsible for that behavior, regardless of whether those genes are located within the body of the particular animal exhibiting that behavior.

J-E Nystrom (2009). Monomorium pharaonis (pharaoh ant), size comparison. Source: Wikimedia Commons. Licensed under CC BY-SA 3.0.

J-E Nystrom (2009). Monomorium pharaonis (pharaoh ant), size comparison. Source: Wikimedia Commons. Licensed under CC BY-SA 3.0.

MEMES IN EVOLUTIONARY BIOLOGY

We commonly know memes as funny images that circulate on Facebook, especially during the Sanremo Festival period, but you may not know that the term “meme” was actually coined by Richard Dawkins. “Meme” comes from the Greek word meaning “imitation,” and it is a term developed on the model of the gene, a genetic unit of heredity capable of replicating itself and producing copies of itself.

The same occurs in mass culture, where an image (or an idea), through imitation, becomes viral and widely known. The term was first used in 1976 in the famous book The Selfish Gene, a work that was both highly appreciated and widely debated, since Dawkins identifies the gene, rather than the individual, as the unit of natural selection (one of the many evolutionary mechanisms that leads to change/evolution). An organism transmits its genes to subsequent generations, and these genes replicate and produce “imitations” of themselves, which are then selected through various evolutionary mechanisms such as natural selection, sexual selection, and others.

Illustration of a meme. Powered by Canva

Illustration of a meme. Powered by Canva

According to Dawkins, cultures can also evolve in a manner analogous to the evolution of populations and living organisms. Many of the ideas transmitted from one generation to the next can increase or decrease the survival chances of the receiving generation, which in turn may pass them on to the next generation. For example, many cultures may develop a design and a method for producing a tool, but the one that develops the most effective methods will have a higher probability of surviving and spreading compared to others. Over time, this idea may be transmitted to a larger portion of the population, which will then adopt those methods. The design of the tool thus acts in a way similar to a biological gene belonging to certain populations and not others, guiding the future of each culture through its presence or absence, or through modifications. Both genes and memes can survive longer than the individual organism that “carries” them. A gene can remain unchanged in the gene pool for hundreds of thousands of years. A useful meme can spread from one individual to another for very long periods after its emergence. Unlike genes, whose success is linked to the survival of the organism’s population, the success of a meme is linked to more subtle factors such as criticism, persuasion, fashion, or group pressure.

Evolution does not only require heredity and natural selection or other evolutionary mechanisms, but also mutation, since memes can also mutate. The ideas that are transmitted can undergo modifications that accumulate over time. I would also add that Dawkins’ idea is based on the fact that memes can propagate independently of their usefulness or functionality, and that they can also sometimes “block” potential cultural transmission between individuals, depending on the context. They can evolve (mutate) very quickly, or become extinct very quickly. In essence, Dawkins argues that natural selection (as an evolutionary mechanism, one of many that “sifts” genes already present within a population) does not only need biological forms for the transmission of information (with the possibility of random variation).

In short, culture evolves like an organism, following the same laws and the same evolutionary mechanisms, but it also has the ability to spread beyond direct relatives.


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