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Red Queen hypothesis

The idea that, in order for a species to maintain a particular niche in an ecosystem and its fitness relative to other species, that species must be constantly undergoing adaptive evolution because the organisms with which it is  coevolving  are themselves undergoing adaptive evolution. When species evolve in accordance with the Red Queen hypothesis, it often results in an evolutionary  arms race .

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What Is the Red Queen Hypothesis?

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Evolution is the changing in species over time. However, with the way ecosystems work on Earth, many species have a close and important relationship with each other to ensure their survival. These symbiotic relationships, such as the predator-prey relationship, keep the biosphere running correctly and keep species from going extinct. This means as one species evolves, it will affect the other species in some way. This coevolution of the species is like an evolutionary arms race that insists that the other species in the relationship must also evolve to survive.

The “Red Queen” hypothesis in evolution is related to the coevolution of species. It states that species must continuously adapt and evolve to pass on genes to the next generation and also to keep from going extinct when other species within a symbiotic relationship are evolving. First proposed in 1973 by Leigh Van Valen, this part of the hypothesis is especially important in a predator-prey relationship or a parasitic relationship.

Predator and Prey

Food sources are arguably one of the most important types of relationships in regards to survival of a species. For instance, if a prey species evolves to become faster over a period of time, the predator needs to adapt and evolve to keep using the prey as a reliable food source. Otherwise, the now faster prey will escape, and the predator will lose a food source and potentially go extinct. However, if the predator becomes faster itself, or evolves in another way like becoming stealthier or a better hunter, then the relationship can continue, and the predators will survive. According to the Red Queen hypothesis, this back and forth coevolution of the species is a constant change with smaller adaptations accumulating over long periods of time.

Sexual Selection

Another part of the Red Queen hypothesis has to do with sexual selection. It relates to the first part of the hypothesis as a mechanism to speed up evolution with the desirable traits. Species that are capable of choosing a mate rather than undergoing asexual reproduction or not having the ability to select a partner can identify characteristics in that partner that are desirable and will produce the more fit offspring for the environment. Hopefully, this mixing of desirable traits will lead to the offspring being chosen through natural selection and the species will continue. This is a particularly helpful mechanism for one species in a symbiotic relationship if the other species cannot undergo sexual selection.

Host and Parasite

An example of this type of interaction would be a host and parasite relationship. Individuals wanting to mate in an area with an abundance of parasitic relationships may be on the lookout for a mate that seems to be immune to the parasite. Since most parasites are asexual or not able to undergo sexual selection, then the species that can choose an immune mate has an evolutionary advantage. The goal would be to produce offspring that have the trait that makes them immune to the parasite. This would make the offspring more fit for the environment and more likely to live long enough to reproduce themselves and pass down the genes.

This hypothesis does not mean that the parasite in this example would not be able to coevolve. There are more ways to accumulate adaptations than just sexual selection of partners. DNA mutations can also produce a change in the gene pool only by chance. All organisms regardless of their reproduction style can have mutations happen at any time. This allows all species, even parasites, to coevolve as the other species in their symbiotic relationships also evolve.

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40 The Red Queen

What does lewis carroll’s red queen have to do with sex.

In this scene from Through the Looking-Glass and What Alice Found There by Lewis Carroll, Alice and the Queen run with all their effort – yet make no progress.  Such is the claim that sex allows organisms to avoid extinction by keeping up in a very odd sort of race.

Alice never could quite make it out, in thinking it over afterwards, how it was that they began: all she remembers is that they were running hand in hand, and the Queen went so fast that it was all she could do to keep up with her: and still the Queen kept crying “Faster! Faster!”, but Alice felt she could not go faster, though she had not breath left to say so. 

The most curious part of the thing was, that the trees and the other things round them never changed their places at all: however fast they went, they never changed their places at all: however fast they went they never seemed to pass anything.  “I wonder if all the things move along with us?” thought poor puzzled Alice.  And the Queen seemed to guess her thoughts, for she cried “Faster! Don’t try to talk!”

Not that Alice had any idea of doing that.  She felt as if she would never be able to talk again, she was getting so much out of breath: and still the Queen cried “Faster! Faster!”, and dragged her along.  “Are we nearly there?” Alice managed to pant out at last.

“Nearly there!” the Queen repeated.  “Why, we passed it ten minutes ago! Faster!” And they ran on for a time in silence, with the wind whistling in Alice’s ears, and almost blowing her hair off her head, she fancied

“Now! Now!” cried the Queen. “Faster! Faster!” And they went so fast that at last they seemed to skim through the air, hardly touching the ground with their feet, till suddenly, just as Alice was getting quite exhausted, they stopped, and she found herself sitting on the ground, breathless and giddy. 

The Queen propped her up against a tree, and said kindly, “You may rest a little, now.”

Alice looked round her in great surprise.  “Why, I do believe we’ve been under this tree the whole time!  Everything’s just as it was!”

“Of course it is, “ said the Queen.  “What would you have it?”

“Well, in our country, “ said Alice, still panting a little, “you’d generally get to somewhere else – if you ran very fast for a long time as we’ve been doing.”

“A slow sort of country!” said the Queen.  “Now here, you see, it takes all the running you can do, to keep in the same place.  If you want to get somewhere else, you must run at least twice as fast as that!”

“I’d rather not try, please!” said Alice….

      Through the Looking-Glass and What Alice Found There by Lewis Carroll

Is sex part of an arms race?

In one human generation, HIV (the virus that causes AIDS) will reproduce over a million times. Given how natural selection works—via heritable variation and differential reproduction—human beings don’t stand a chance against this virus. How can we possibly adapt to such a fast-moving target? For that matter, how can any longer-lived organism compete with a quickly reproducing and quickly evolving enemy? Many of these enemies, or pathogens , such as viruses and bacteria, are also numerous and difficult to detect—invisible to the naked eye, they can enter a host’s body silently and reproduce with a fervor while their victims remain blissfully unaware. Given these challenges, how can any host organism defend itself against its would-be attackers? According to one hypothesis, outwitting pathogens is the whole point of sex.

The Red Queen

We are in the midst of an evolutionary arms race , in which host and parasitic pathogen must constantly adapt. Parasites must adapt to the host’s natural defenses, and host populations are under pressure to keep up with their ever-changing parasites.  This reciprocal evolution between two types of organisms (in this case, host and parasite) is a type of coevolution. According to the Red Queen Hypothesis , sex exists as a mechanism for keeping up with rapidly coevolving pathogens. By generating genetic diversity, sex makes host organisms a moving target.  Like Alice and the Red Queen in Lewis Carroll’s novel (Box 3), both host and parasite are running a race in which neither makes any observable progress. Yet, if the host organisms didn’t change dramatically with each new generation (if they didn’t have sex), they might go extinct.

Parasites adapt to exploit the most common type of host.  Therefore, a host that can produce offspring that have novel defenses against parasites would have an advantage over an organism producing clones–simply by making offspring that are different.

Introductory Biology: Evolutionary and Ecological Perspectives Copyright © by Various Authors - See Each Chapter Attribution is licensed under a Creative Commons Attribution 4.0 International License , except where otherwise noted.

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The Red Queen was right: Life must continually evolve to avoid extinction

Biologists quote Lewis Carroll when arguing that survival is a constant struggle to adapt and evolve. But is it true, or do groups die out because they experience a run of bad luck? UC Berkeley biologists tested these hypotheses and found that it’s not luck at all, but failure to adapt to a deteriorating environment, that leads to extinction.

By Robert Sanders

June 20, 2013

The death of individual species shouldn’t be the only concern for biologists worried about animal groups, such as frogs or the “big cats,” going extinct. A University of California, Berkeley, study has found that a lack of new, emerging species also contributes to extinction.

As the Red Queen told Alice, “it takes all the running you can do, to keep in the same place.” Similarly, animals and plants must continually adapt and evolve just to avoid going extinct. (Illustration by Sir John Tenniel from Lewis Carroll’s “Through the Looking-Glass,” 1871)

“Virtually no biologist thinks about the failure to originate as being a major factor in the long term causes of extinction,” said Charles Marshall, director of the UC Berkeley Museum of Paleontology and professor of integrative biology, and co-author of the report. “But we found that a decrease in the origin of new species is just as important as increased extinction rate in driving mammals to extinction.”

The effects of such a decrease would play out over millions of years, Marshall said, not rapidly, like the global change Earth is experiencing from human activities. Yet, the findings should help biologists understand the pressures on today’s flora and fauna and what drove evolution and extinction in the past, he added.

The results, published June 20 in the journal Science Express , come from a study of 19 groups of mammals that either are extinct or, in the case of horses, elephants, rhinos and others, are in decline from a past peak in diversity. All are richly represented in the fossil record and had their origins sometime in the last 66 million years, during the Cenozoic Era.

The study was designed to test a popular evolutionary theory called the Red Queen hypothesis, named after Lewis Carroll’s character who, in the book “Through the Looking Glass,” described her country as a place where “it takes all the running you can do, to keep in the same place.”

In biology, this means that animals and plants don’t just disappear because of bad luck in a static and unchanging environment, like a gambler losing it all to a run of bad luck at the slot machines. Instead, they face constant change – a deteriorating environment and more successful competitors and predators – that requires them to continually adapt and evolve new species just to survive.

Though the specific cause of declining originations and rising extinctions for these groups is unclear, the researchers concluded that the mammals’ death was not just dumb luck.

“Each group has either lost, or is losing, to an increasingly difficult environment,” Marshall said. “These groups’ demise was at least in part due to loss to the Red Queen – that is, a failure to keep pace with a deteriorating environment.”

Marshall and former UC Berkeley post-doctoral fellow Tiago Quental found that the animal groups were initially driven to higher diversity until they reached the carrying capacity of their environment, or the maximum number of species their environment could hold. After that, their environment deteriorated to the point where there was too much diversity to be sustained, leading to their extinction.

“In fact, our data suggest that biological systems may never be in equilibrium at all, with groups expanding and contracting under persistent and rather, geologically speaking, rapid change,” he said.

Marshall and Quental, who is now at the University of Sao Paolo, Brazil, will present their results in two talks this Saturday, June 22, at the Evolution 2013 meeting in Snowbird, Utah.

MORE INFORMATION

• How the Red Queen drives terrestrial mammals to extinction ( Science Express )
• Charles Marshall’s laboratory web site
• University of California Museum of Paleontology

Red Queen Hypothesis

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The Red Queen hypothesis was proposed by Van Valen in 1993, in a famous paper entitled “A New Evolutionary Law.” On the basis of a detailed study of the extinction rates of species during geologic times, Van Valen suggested that any adaptation of a species modifies the environment of the other species living with it and constrains them to new adaptations, because of the limitation of resources: species play a “null sum game.” This causes a runaway process which is the cause of the indefinite complexification of species. Van Valen used the term “Red Queen hypothesis” by reference to the novel of Lewis Carrol where Alice and the Red Queen run “to keep in the same place.”

It is probably in the world of parasites that the Red Queen hypothesis has been most illustrated and discussed.

A first reason why parasites have something to do with Van Valen’s hypothesis is that when a parasite species and a host species coevolve for a long time, each one exerts selective pressures on the other. Thus,...

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Institut für Zoomorphologie, Zellbiologie und Parasitologie, Heinrich-Heine-Universität, Universitätsstraβe 1, 40225, Düsseldorf, Germany

Heinz Mehlhorn Dr. ( Professor ) ( Professor )

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(2008). Red Queen Hypothesis. In: Mehlhorn, H. (eds) Encyclopedia of Parasitology. Springer, Berlin, Heidelberg. https://doi.org/10.1007/978-3-540-48996-2_2666

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• v.14(5); 2018 May

Getting somewhere with the Red Queen: chasing a biologically modern definition of the hypothesis

Luke c. strotz.

1 Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, KS 66045, USA

2 Biodiversity Institute, University of Kansas, Lawrence, KS 66045, USA

Marianna Simões

Matthew g. girard, laura breitkreuz, julien kimmig, bruce s. lieberman, associated data.

This article has no additional data.

The Red Queen hypothesis (RQH) is both familiar and murky, with a scope and range that has broadened beyond its original focus. Although originally developed in the palaeontological arena, it now encompasses many evolutionary theories that champion biotic interactions as significant mechanisms for evolutionary change. As such it de-emphasizes the important role of abiotic drivers in evolution, even though such a role is frequently posited to be pivotal. Concomitant with this shift in focus, several studies challenged the validity of the RQH and downplayed its propriety. Herein, we examine in detail the assumptions that underpin the RQH in the hopes of furthering conceptual understanding and promoting appropriate application of the hypothesis. We identify issues and inconsistencies with the assumptions of the RQH, and propose a redefinition where the Red Queen's reign is restricted to certain types of biotic interactions and evolutionary patterns occurring at the population level.

1. ‘Down the rabbit hole’ 1 : introduction

The Red Queen hypothesis (RQH) was first proposed by Van Valen [ 1 ] to explain a pattern he argued was manifest in the fossil record involving component members of several major taxonomic groups: survivorship curves that were linear when plotted against geologic time. The RQH contains several additional elements Van Valen [ 1 ] derived from this pattern. First, in any taxonomic group that occupies the same adaptive landscape, the probability of survival is independent of age throughout its existence. Then, Van Valen [ 1 ] took this interpretation one step further and concluded that all members of such groups had an equal probability of extinction. This aspect of the RQH he termed the ‘Law of Constant Extinction’ which was held to be applicable across different organizational (e.g. population, community), and taxonomic (e.g. species, genera, families) levels. Finally, Van Valen [ 1 ] suggested that the RQH involved omnipresent competitive interactions among taxonomic groups; these were continually changing, but they were not getting relatively better in a competitive sense through time such that there was a zero-sum expectation ( figure 1 ). Instead, they were metaphorically running in place and not getting anywhere: like the eponymous Red Queen from Lewis Carroll's ‘Through the Looking-Glass, and What Alice Found There’.

Evolutionary change under Red Queen hypothesis-type dynamics versus Court Jester hypothesis-type dynamics. The blue line represents the abiotic environment. Species A (green) represents a potential prey organism. Species B (purple) represents its potential predator. Species C (black) also preys on species A. Species D (orange) is a descendant taxon of species B. In both scenarios (Red Queen and Court Jester), species B goes extinct (represented by the dotted line). Under the Red Queen scenario ( sensu Van Valen [ 1 ]), the extinction of species B is due to species A shifting from adaptive zone 1 to adaptive zone 2 as it moves towards a fitness optimum and ultimately exceeds the relevant traits of species B (traits relevant to the capacity of species B to capture and consume species A). The new adaptive zone that species A now occupies contains a different predator species (species C) whose fitness is affected by the arrival of species A, as per Van Valen's [ 1 ] zero-sum assumption. In this scenario, abiotic parameters remain unchanged and evolutionary change can still occur. In the Court Jester scenario, the extinction of species B is due to environmental changes that result in suboptimal conditions for species B. Populations within the species become isolated from one another, population sizes decrease, and almost all of the component populations die off, such that species B goes extinct; however, in one case an isolated population of species B diverges, survives and becomes a new species (species D). No changes in the adaptive zone of species A occur in this scenario, nor did they the cause the extinction of species B. Both hypotheses have different evolutionary and spatial scales. The RQH operates across individual populations on small spatial and short temporal scales, leading to differential survival of populations within communities. By contrast, the Court Jester hypothesis is tied to large-scale shifts in the physical environment, which would affect multifarious populations from species in different clades, with each population potentially responding individualistically to the perturbation [ 2 ].

Many aspects of Van Valen's findings have stimulated extensive debate and discussion. For instance, his purported link between extinction probability and species age has been disputed [ 3 – 7 ]. The RQH has also been the subject of reviews (e.g. [ 8 – 10 ]) and a variety of modelling-based studies (e.g. [ 11 – 15 ]). Some of these endorsed aspects of the core predictions of the RQH, others challenged them. Moreover, additional definitions of the RQH have been proposed [ 10 ] that differ from Van Valen [ 1 ]. Scientists have also attempted to use the RQH to explain phenomena beyond its original purview, for example, the dynamics of some host–parasite systems [ 16 ] and the role that these coevolutionary relationships may play in the maintenance of sexual reproduction [ 8 , 17 – 19 ].

Even as the scope of the RQH has broadened, at its core the RQH retains a key element: the primary drivers of macroevolution are held to be biotic interactions, in particular, the effects that the origin of a new trait in one group (populations, species, etc.) has on each group it interacts with [ 9 ]. This feature is not exclusive to the RQH. Further, there is an emphasis in the RQH on groups interacting with equal effect [ 1 ]. This is potentially problematic, as real ecological dynamics are usually far more complicated [ 20 ] and groups rarely have equal effects on each other [ 21 ].

Also, notably many studies [ 2 , 3 , 20 – 24 ] have disputed the validity of numerous aspects of Van Valen's [ 1 ] original RQH. Thus, it is perhaps surprising that the RQH continues to receive support and evolve as a concept [ 9 , 25 – 28 ]. In the light of these theoretical and conceptual peregrinations, we re-examine the RQH, discuss original and subsequent expositions and put forward a single definition, informed by Van Valen's [ 1 ] original exposition, that also accounts for subsequent treatments. The aim is to provide clarity to what has at times been a murky topic in evolutionary biology.

2. ‘It's no use going back to yesterday, because I was a different person then’: the RQH evolves

Stenseth & Maynard Smith [ 12 ] suggested rejecting the RQH's zero-sum expectation and proposed that RQH dynamics may only apply in ecosystems where evolutionary rates are greater than zero, where evolution is mediated by biotic interactions, and where the physical environment remains unchanged. The purpose of this was not to refute the RQH, but to provide the RQH with an alternate null hypothesis where environmental change is the impediment to evolutionary stasis, and evolutionary advances by one species need not necessarily result in a net negative effect of the same magnitude across other species. While this new hypothesis was not the RQH sensu Van Valen [ 1 ], it did open up the application of a RQH-like framework beyond its original domain and began a trend of placing different evolutionary phenomena under the banner of the RQH that did not entirely align with the RQH sensu Van Valen [ 1 ]. For instance, it is suggested that the RQH provides a mechanism for the evolution and maintenance of sex by explaining the value of recombination due to the negative frequency-dependent selection associated with parasitism [ 8 , 17 , 18 ], with sexual lineages better at evading parasites via genetic recombination, thereby forcing continual coevolution on the part of the parasite to maintain a constant level of virulence [ 16 ]. Similar types of continual coevolutionary patterns have also been proposed for predator–prey relationships [ 19 ], with predators maintaining sexual reproduction to preserve a constant level of predation success as variable prey populations within species shift in abundance between those with a greater capacity for obtaining nutrients and those better able to defend against predation.

In an attempt to clarify and categorize the growing body of RQH-influenced ideas, an important synthesis was provided by Brockhurst et al . [ 10 ], who argued that there were three distinct classes of concepts that aligned with the RQH based on the patterns they displayed and the processes involved. The work of Brockhurst et al . [ 10 ] also continued the trend of the RQH evolving beyond the original purview of Van Valen [ 1 ]. Brockhurst et al .'s [ 10 ] classes were designated the Escalatory, Fluctuating and Chase. The Escalatory-RQH occurs when species interactions lead to directional selection and both interacting species move towards a fitness optimum as each struggles to ‘exceed’ the relevant trait of the other species [ 10 ]. Such a dynamic is posited in the case of evolutionary arms races. The Fluctuating-RQH is associated with an oscillating mode of selection, where two antagonist species oscillate backwards and forwards between fitness optima, with one interactor always lagging behind the other [ 10 ]. This involves continual, yet non-directional, evolutionary motion for both antagonists, analogous to constrained stasis or a random walk in species morphology that produces no net change over the long term [ 29 , 30 ]. Finally, the Chase-RQH supposes that across the range of two interacting or co-evolving species, respective populations may be responding in different ways to the biotic milieu they experience [ 10 ]. As populations of the chased antagonist seek to escape co-occurring populations of the chaser through the evolution of novelty, diversity within populations becomes reduced but divergence between populations increases as they spread across a multidimensional phenotypic space. All three classes outlined by Brockhurst et al . [ 10 ] invoke biotic interactions among two groups as the drivers of evolutionary change. The Escalatory-RQH approximates the RQH sensu Van Valen [ 1 ]: a key difference is the latter focuses on higher taxa. The Chase-RQH, however, diverges from the RQH sensu Van Valen [ 1 ] because it involves several interacting component populations of different species, each evolving in varying directions due to distinct selective pressures. The Fluctuating-RQH also potentially diverges from the RQH sensu Van Valen [ 1 ] if the emphasis is placed on changes in specific fitness or phenotypic states, because species are hopping back and forth between distinct states rather than continually running in place. Alternatively, it is possible that if the change in question is migration across the evolutionary landscape, or changes in the dynamics of species interactions, then the relevant species are indeed running in place and the fluctuating-RQH can be considered equivalent to the RQH sensu Van Valen [ 1 ].

3. ‘Off with her head!’: problematic aspects of the RQH

Acceptance of the RQH has not been universal, and a number of authors have either implicitly or explicitly opposed Van Valen's [ 1 ] conclusions. For instance, the competitive species interactions invoked by the RQH have been shown as unlikely to result in persistent evolutionary change [ 31 ]. Questions have also been raised as to whether the taxonomic survivorship curves presented by Van Valen [ 1 ] are truly linear [ 5 , 7 , 32 – 35 ]. While evidence from planktic microfossils has been used to support log-linearity of species-level survivorship curves [ 11 , 36 – 39 ], results for planktic foraminifera have, by contrast, demonstrated a positive relationship between extinction risk and species age [ 6 , 40 , 41 ]. Mass extinction has been singled out as one significant phenomenon that causes groups to deviate from constant extinction over time (e.g. [ 35 , 42 ]). Van Valen [ 1 ] himself noted that mass extinctions in specific clades (e.g. ammonites) were exceptions to the ‘Law of Constant Extinction’ as they represent times of exceptional elevation of extinction rates. Intriguingly though, if mass extinctions truly eliminate large numbers of species effectively at random then, under certain circumstances of prior diversification, they could result in situations where the probability of extinction of species is independent of its duration [ 3 ].

Conceptual criticisms of the RQH have also focused on whether, even assuming taxonomic survivorship curves are linear, the ‘new evolutionary law’ Van Valen [ 1 ] erected was valid [ 3 , 4 , 43 , 44 ]. For example, McCune [ 3 ] concluded that, while the probability of extinction of taxa within a clade may be constant with respect to the duration of those taxa, this does not mean that the rate or the probability of extinction is constant per unit time. She thus argued that the RQH is only one of many potential phenomena that might explain linear taxonomic survivorship curves.

The RQH also depends upon substantive phyletic speciation and associated pseudoextinction [ 4 , 12 ]. This creates a paradox for the RQH because, if phyletic speciation is a primary evolutionary mode, this means that the rate of extinction will be directly correlated with the rate of evolution. Yet the RQH posits that species extinction should be independent of duration. As Vrba [ 4 ] recognized, the rate of phyletic speciation cannot be independent of itself.

4. ‘I'm not crazy. My reality is just different than yours’: abiotic alternatives to the RQH

Another significant criticism of the RQH stems from the limited role it imputes to abiotic factors as important drivers of evolutionary change [ 2 , 4 , 24 ]. Evidence from a variety of sources [ 45 – 51 ] uphold abiotic factors as important drivers of evolution and speciation. This has led to proposed alternatives to the RQH which focus on the physical environment as the main driver of evolution. The most prominent of these is the ‘Court Jester’ hypothesis [ 23 ] ( figure 1 ), with the name chosen to highlight the capricious effects environmental changes can have on evolution. This is in contradiction to the more predictable effects that might be associated with the RQH. The Court Jester attempts to unite under one concept the plethora of previously proposed ideas that support abiotic factors as main drivers of evolutionary change (e.g. ‘turnover-pulse hypothesis’ [ 4 ]; ‘stationary model’ [ 12 ] and ‘coordinated stasis’ [ 52 ]).

Proponents of abiotic change as the chief driver of evolution have been particularly critical of the assertion that competition between groups at taxonomic ranks higher than the species, where the RQH sensu Van Valen [ 1 ] is focused, could result in these groups diverging or going extinct [ 2 , 4 , 24 ]. If and when the RQH does operate, it should be at the level of individual populations at small spatial and short temporal scales. Entities at higher hierarchical levels ( sensu [ 2 , 4 ]), such as clades which consist of many species, should not be expected to respond as a unit ([ 45 ]; though see [ 53 ] for a divergent opinion). Indeed, there is scant evidence that the RQH operates at scales involving entire continents or millions of years [ 2 , 4 , 23 , 24 ]. Instead, individual populations of species living in different communities and climates would interact in different ways with numerous populations of other species across the totality of their ranges, and respond individualistically to any perturbations [ 2 , 4 , 23 , 24 ]. While refocusing the RQH onto the species, and especially the population level, would address these concerns [ 9 ], it is not the same as the RQH sensu Van Valen [ 1 ].

5. ‘Who in the world am I? Ah, that's the great puzzle’: what is the proper domain of the RQH?

As the RQH has been increasingly applied beyond Van Valen's [ 1 ] original focus, it has become increasingly difficult to evaluate its legitimacy. It is first and foremost necessary to ascertain whether the RQH sensu Van Valen [ 1 ] has been generally upheld. If it has been, then the RQH provides an explanation for how biotic interactions could drive phenotypic change, even if only under certain circumstances. However, if it is not generally upheld by most studies, then the question becomes which parts of the RQH should be retained in evolutionary theory and how should the RQH be viewed in the future?

It has been proposed that rejecting the RQH is only possible by demonstrating that evolutionary and ecological changes of organisms (presumably at the species or population level) are primarily due to abiotic change, while at the same time also considering the effect of biotic interactions or abiotic change that is biotically driven [ 9 ]. We diverge from this proposal as this set of conditions is probably unachievable, because a period of constancy of any potential abiotic factors is virtually absent from the geological record [ 14 ]. Another challenge to evaluating the RQH is that biotic and abiotic factors can interact to drive macroevolution [ 54 , 55 ], making it hard to differentiate primary biotically driven evolution from secondary biotically driven evolution instigated by abiotic forcing. Because of these challenges, here we focus on an alternative method to assess the validity of the RQH sensu Van Valen [ 1 ]. As would be the case for any hypothesis, if any (or several) of the core assumptions of the RQH are found to be spurious, then the hypothesis itself would be difficult to uphold, and usage should only be done with significant caution and caveats.

Upon consideration of the evidence for and against the RQH sensu Van Valen [ 1 ] as a valid macroevolutionary concept, potential problems with the hypothesis emerge. Of greatest concern are the evidence-based [ 5 – 7 , 32 – 35 ] and concept-based [ 3 , 4 ] refutations of the foundational assumption of the RQH that taxonomic survivorship curves are linear. This undercuts the very notion of Van Valen's [ 1 ] ‘Law of Constant Extinction’. These refutations are supported by an even larger body of the literature in palaeontology and community ecology, demonstrating that extinction is associated with a range of parameters that are not purely stochastic (e.g. geographical range size is a demonstrated key predictor of extinction [ 55 ]). Van Valen, despite erecting his ‘law’, acknowledged ‘the probability of extinction is not constant over geological time’ ([ 1 ], p. 18) and that constant extinction only prevails if extinctions associated with major perturbations are ignored. Mass extinctions, of course, are a significant feature of the history of life. If the assumption of constant extinction rates is invalid, and extinction probabilities are not age independent, this impugns the RQH as an explanation of macroevolutionary patterns. Evolutionary advances by one species need not produce a net negative effect of the same magnitude across all other coexisting species, because no mechanism is required to maintain equal extinction probabilities. The apparent difficulty of reducing macroevolutionary dynamics to a zero-sum process [ 12 ] finds meaning when the ‘Law of Constant Extinction’ is rejected [ 20 , 21 ].

There are additional issues that, when considered singularly, would not be enough to refute the foundational assumptions of the RQH but, when considered collectively, call into question their cogency. First, Van Valen [ 1 ] used data from groups that have subsequently been identified as paraphyletic, such that they lack evolutionary significance [ 56 ]. Second, only five of the 56 clades Van Valen [ 1 ] analysed were treated at the species level, with the remaining majority comprising either genera or families. The notion that competition, selection or anagenesis could involve higher taxa is inconsistent with basic evolutionary principles. For example, genera are arbitrary units that are not necessarily monophyletic or equivalent across clades [ 56 ] and it is unlikely evolutionary processes can be applied to them.

A further difficulty with upholding the RQH involves the phenomenon of pseudoextinction. Although Van Valen [ 1 ] noted that pseudoextinction is an infrequent process in the fossil record (at higher taxonomic levels), frequent pseudoextinction is perforce necessary in any system where the RQH is the explanatory mechanism for evolutionary change [ 4 , 12 , 57 ]. Like constant extinction, frequent pseudoextinction has been refuted in the literature [ 58 ], and does not make sense in light of modern phylogenetic understanding. Analyses of species origination for planktic foraminifera, a group where taxon durations can be accurately estimated at the species level, have demonstrated pseudoextinction to be a rare occurrence (less than 10%) at the macroevolutionary scale [ 58 , 59 ], and once putative ‘archetypal’ examples of anagenesis with concomitant pseudoextinction have subsequently been shown to involve cladogenesis [ 60 ].

Ultimately, given the challenges to the evidence and core assumptions underlying Van Valen's [ 1 ] RQH, it seems hard to advocate that his microevolutionary mechanism of intrinsic biotic conflicts is what drives the macroevolutionary trends observed in the fossil record. There may still be a place for a RQH-like framework but, if so, it operates at the level of populations within ecosystems [ 2 , 4 , 24 ] and is not the RQH sensu Van Valen [ 1 ].

6. ‘Everything's got a moral, if only you can find it’: a contemporary definition of the RQH

The paucity of support for the RQH sensu Van Valen [ 1 ] does not mean that we propose no conditions exist where biotic interactions could be a significant mechanism for evolutionary change. It also does not take away from the fact that Van Valen's [ 1 ] original RQH was highly valuable, stimulated a variety of important research and greatly furthered conceptual understanding. Antagonistic interactions are potential examples where a qualified version of the RQH could conceivably apply, such as populations of hosts and their parasites or predators and their prey [ 19 , 61 ].

We propose that were one to take Van Valen's RQH and modify it based on developments in evolutionary theory, palaeontology and phylogenetics made after 1973, then what results is Brockhurst et al .'s [ 10 ] Chase-RQH. Although Brockhurst et al . [ 10 ] did not go so far as to nominate the Chase-RQH (or any of their other classes of Red Queen) as a replacement of the RQH sensu Van Valen [ 1 ], we do so here ( figure 2 ). It is not Van Valen's [ 1 ] RQH because it does not focus on constant extinction rates within higher taxonomic groups, nor does it claim that the probability of extinction of any taxonomic group is independent of its duration, but it does capture coevolutionary relationships where two interacting, antagonistic populations are continually changing yet neither is ‘improving’ relative to the other. It also parallels Thompson's [ 62 ] strongly supported view of coevolution, which emphasized the geographical mosaic of the phenomenon. Further, it agrees with Eldredge's [ 2 ], Vrba's [ 4 ], Barnosky's [ 23 ], Benton's [ 24 ] and Liow et al .'s [ 9 ] contention that the RQH be refocused at the species/population level and that Red Queen phenomena occur at the level of populations within ecosystems, not species or higher taxa across vast tracts of geographical space. Brockhurst et al .'s [ 10 ] Chase-RQH also resolves the problems with Van Valen's [ 1 ] RQH that pertain to pseudoextinction. In fact, the Chase-RQH can lead to cladogenesis, as individual populations may deviate from the species as a whole ( figure 2 ), such that a descendant may evolve while its ancestor persists. This divergence among populations subject to different selection regimes places Chase-RQH within the tenets of modern microevolutionary theory. The Chase-RQH also aligns with models linking host–parasite interactions to the evolution and maintenance of sex. Most importantly, the Chase-RQH establishes a link between microevolutionary processes and macroevolutionary patterns that is internally consistent with current thinking in evolutionary biology.

A contemporary definition of RQH based upon Brockhurst et al .'s [ 10 ] Chase Red Queen and informed by Vrba's [ 4 ] turnover pulse hypothesis and Barnosky's [ 23 ] Court Jester hypothesis. Two interacting co-evolving species (species A and B) have seemingly fixed relative fitness over time at the macroscale. A detailed examination (first box) shows at the microscale that actually species A is a chased antagonist pursued by species B across an adaptive zone, with a lag in the response of species B relative to the shifts in mean fitness of species A. These shifts in mean fitness for species A and B represent disparate reactions by individual populations of each species as they respond individualistically to biotic or abiotic perturbations (second box). As populations of the chaser seek to keep pace with co-occurring populations of the chased antagonist, individual populations deviate from the species mean trait values and may even go extinct (e.g. species B, pop 4). Because the mean relative fitness of the two species is constant over time, the relative fitness of co-occurring populations of species A versus species B must be maintained over time, but the relative fitness of non-co-occurring populations of species A versus species B need not remain static. Significant abiotic events can potentially alter this balance, with individual populations of either species becoming isolated and forming a new species, or the extinction of key populations leading to the overall extinction of either species (but not necessarily both).

Acknowledgements

We thank Mabel Alvarado, Rebecca Dorward, Jennifer C. Giron Duque, Kaylee Herzog, Kayla Kolis and Ryan Ridder for their contributions to the preparation of this manuscript. We would also like to thank Paul Craze, the handling editor and three anonymous reviewers whose comments helped to improve our paper.

1 In the tradition of Leigh Van Valen and inspired by his choice of Alice's encounter with the Red Queen as an apt allegory for biotically driven evolutionary dynamics, we use relevant quotes from Lewis Caroll's ‘Alice's Adventures in Wonderland’ or ‘Through the Looking-Glass, and What Alice Found There’ as subheadings for each of our sections.

Data accessibility

Competing interests.

We declare we have no competing interests

This research was supported by NSF ADBC-1206757 and DBI-1602067.

This year's Darwin Review: Revisiting the Red Queen

The red queen hypothesis states that in the battle for resources, species must continuously evolve just to keep up with their enemies, who themselves also evolve in response. this year, we revisit the seminal theory..

Proceedings B Darwin Reviews are special reviews published up to once a year. The aim of the Darwin Review is to showcase ideas and/or a field in biological science that is of very high interest to the whole diverse readership of Proceedings B, often being of particular relevance to strategic growth areas, importance to policy makers and/or having a bearing on the public that fund our science.

This year our Darwin review revisits a seminal theory in evolutionary research, Van Vaalen’s Red Queen Hypothesis. 40 years after its initial proposal the Red Queen is still informing research. Here the authors discuss their review and why now was the right time to highlight the Red Queen’s enduring legacy. You can read the full article here . Our previous Darwin review was on evolutionary medicine and can be found here .

Tell us about the Red Queen hypothesis.

The Red Queen hypothesis was proposed over 40 years ago by the late evolutionary biologist Leigh Van Valen. It advanced evolutionary thinking beyond the idea that organisms were merely matched to their physical environment by suggesting that interactions between species (such as between hosts and parasites, predators and prey) would also be important in driving evolutionary change. In essence, the Red Queen hypothesis states that in the battle for resources, species must continuously evolve just to keep up with their enemies, who themselves also evolve in response. The result is that species constantly change but, relative to their enemies, don’t actually get any fitter – like running on an evolutionary treadmill. The name for the theory came from Lewis Carroll’s ‘Through the Looking Glass’ (aka Alice in Wonderland). Alice finds herself in a race with the Red Queen, and despite running as fast as she can, Alice stays in the same place. The Red Queen hypothesis, doubtless partly due to this imaginative metaphor, has become one of the most influential ideas in evolution.

How has the theory influenced evolutionary biology research since its original proposal?

Van Valen was interested in macroevolution, that is speciation and extinction, and used the Red Queen hypothesis to explain the apparently constant rates of extinction observed in the fossil record. And although this pattern has subsequently been challenged, Red Queen thinking is still important in macroevolution in terms of understanding the biological, as opposed to environmental, causes of diversification and extinction. In the 1980s, the utility of the Red Queen concept was rediscovered as a way of explaining the ubiquity of sexual reproduction in nature. The idea starts by considering that hosts and parasites are evolving together. If the hosts are sexual females, they can produce offspring that are genetically diverse and therefore avoid parasite infection. However, if hosts are clonal females, their offspring lack the ability to be as genetically diverse and will be more susceptible to attack by parasites. Thus, coevolution with parasites prevents the clones from taking over and gives sexual species an advantage. Increasingly, biologists think of the Red Queen hypothesis, not only as an explanation for sex, but also as a means of explaining rapid evolutionary change in hosts and their parasites in general.

What prompted you to write this review?

The adoption of the Red Queen as an explanation for the evolution and widespread maintenance of sex led to a narrowing of the definition of the Red Queen hypothesis. This meant that the far richer perspective of Van Valen’s original conceptual leap was in danger of being lost. To some extent, we wrote the review because we believed it was necessary to reemphasize a broader importance of the Red Queen hypothesis because it offers a powerful way in which to understand natural communities, how they evolve, and how they work. It is an important and useful idea. We were also inspired by the data emerging from new approaches of studying species interactions, including genome sequencing and experimental evolution in the laboratory. These present a more complex and nuanced picture of the effect of species interactions on evolution that is entirely consistent with a Red Queen view of nature. Additionally, we were aware that much of theory developed to understand how conflicts between species evolve could be usefully applied to understand conflicts within species. A variety of conflicts have been defined in this context: intragenomic conflicts (e.g. over the representation of chromosomes in gametes during meiosis), sexual conflicts (e.g. between males and females over reproductive effort and timing) and parent / offspring conflicts (e.g. over the allocation of resources to progeny). However, the evolutionary dynamics of these conflicts has never formally been synthesized within the Red Queen concept, despite the clear ability for these conflicts to generate the type of ‘running to stay still’ evolutionary dynamics. One of our aims in this review was therefore also to attempt this synthesis.

What does the future hold the red queen hypothesis?

It will certainly be informative to see how useful and explanatory is the incorporation of conflicts within species into the Red Queen framework. Beyond this, we recognise that much of our appreciation of the Red Queen currently comes from studying binary relationships (one parasite in one host). Extensions of this to more realistic scenarios, by incorporating parasites that evolve with a range of hosts, and hosts that evolve with a range of parasites, are therefore needed. How these will affect the speed of the Red Queen needs to be determined. Finally, there has been increasing use of comparative genomics to examine evolutionary dynamics. However, these analyses are limited in that the function of many genes is not known, making it difficult to determine the contribution of the Red Queen. Commonly, for instance, expression patterns indicate that specific genes are involved in male-female interactions, and could therefore be subject to coevolution. However, at the moment, we cannot differentiate the subset of genes that are involved with the male-female interface that are likely to be subject to Red Queen forces from those involved in a variety of other sex-specific physiological functions. We also have incomplete ascertainment of genes at the host-pathogen boundary: we understand host genes that are generically involved in defence much better than those which alter microbial invasion into a cell. We expect the results of comparative genomic analysis of Red Queen type processes to be made sharper as our understanding of gene function improves.

Meet the authors

Michael Brockhurst is Professor of Evolutionary Biology and a 50th Anniversary Chair at the University of York. He uses experimental evolution approaches to study the coevolution of species interactions with a particular focus on bacteria and their viral and genetic parasites. He is interested in the applied consequences of rapid microbial evolution in natural communities especially in clinically important pathogenic microbes.

Greg Hurst is Professor of Evolutionary Biology at the University of Liverpool. His main goal is to determine the ecological and evolutionary importance of heritable symbionts of insects (where symbiont is broadly defined and includes parasitism), and has an interest intragenomic and intra-specific conflicts more generally. His study animals include Drosophila, Nasonia wasps, butterflies and ladybirds, and the symbionts studied are largely microbial.

Kayla King is an Associate Professor at the University of Oxford. Her research explores the ecology and evolution of species interactions to ask fundamental questions about the maintenance of genetic and community-level diversity, sexual reproduction, and rapid evolutionary change. She focuses on interactions between hosts and their parasites as well as their microbiota.

Judith Mank is Professor of Evolutionary and Comparative Biology at University College London. She is interested in the constraints imposed on the genome by sexual conflict, and how selection navigates these restrictions of genome architecture to create intra-specific phenotypic diversity in the form of sexual dimorphism. She works on a range of study organisms, most recently birds, fish and flies.

Steve Paterson is Professor of Evolutionary Biology and a director of The Centre for Genomic Research at the University of Liverpool. He is primarily interested in understanding the forces that shape genetic diversity within host and parasite genomes. New genomic methods that allow us to address this question make it clear how much we owe to the Red Queen for our understanding of genome evolution and of the novel biology yet to be discovered from biotic interactions.

Tracey Chapman is Professor of Evolutionary Biology in the School of Biological Sciences at the University of East Anglia. She is interested in understanding the nature and the evolutionary potential of reproductive interactions between males and females. Such interactions are often subject to sexual conflict and can generate rapid evolutionary change.

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EvolutionBiology.com

Red queen principle.

Through the Looking Glass – Lewis Carroll

The Red Queen Hypothesis in biology states that species continually need to change to keep up with the competition. If a species would stop changing, it would lose the competition with the other species that do continue to change. If you take for example the relationship between a parasite and its host. Both the parasite and the host are involved in an arms race with each other. There is pressure on the host to evolve to become resistant to the parasite and there is pressure on the parasite to evolve ways to cope with the resistance of the host. Both species need to change genetically to keep up with the changes in the other species.

The Red Queen Principle is an important theory because it is used in explaining sexual reproduction , the importance of genetic diversity and the speed of evolution . From the Red Queen Principe follows that species are never “finished”, extinction probability does not increase with existence age of the species and the speed of genetic change over time is important for evolution and survival of species. Species with a quicker generation time will have the ability to evolve faster , giving them an advantage in an arms race. This calls for a mechanism to increase speed of evolution. Sexual reproduction is one way to increase evolution speed in a species, because it allows for new mutations to spread fast in the population and new combinations of alleles to occur faster. It is thought that species with long generation time have to have sexual reproduction to be able to stay in the race with species with a short generation time.

The Red Queen Hypothesis was formulated in 1973 by Leigh Van Valen.

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• Published: 24 February 2010

Antagonistic coevolution accelerates molecular evolution

• Steve Paterson 1   na1 ,
• Tom Vogwill 1   na1 ,
• Angus Buckling 2 ,
• Rebecca Benmayor 2 ,
• Andrew J. Spiers 3 ,
• Nicholas R. Thomson 4 ,
• Mike Quail 4 ,
• Frances Smith 4 ,
• Danielle Walker 4 ,
• Ben Libberton 1 ,
• Andrew Fenton 1 ,
• Neil Hall 1 &
• Michael A. Brockhurst 1   na1  

Nature volume  464 ,  pages 275–278 ( 2010 ) Cite this article

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• Coevolution
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The Red Queen hypothesis proposes that coevolution of interacting species (such as hosts and parasites) should drive molecular evolution through continual natural selection for adaptation and counter-adaptation 1 , 2 , 3 . Although the divergence observed at some host-resistance 4 , 5 , 6 and parasite-infectivity 7 , 8 , 9 genes is consistent with this, the long time periods typically required to study coevolution have so far prevented any direct empirical test. Here we show, using experimental populations of the bacterium Pseudomonas fluorescens SBW25 and its viral parasite, phage Φ2 (refs 10 , 11 ), that the rate of molecular evolution in the phage was far higher when both bacterium and phage coevolved with each other than when phage evolved against a constant host genotype. Coevolution also resulted in far greater genetic divergence between replicate populations, which was correlated with the range of hosts that coevolved phage were able to infect. Consistent with this, the most rapidly evolving phage genes under coevolution were those involved in host infection. These results demonstrate, at both the genomic and phenotypic level, that antagonistic coevolution is a cause of rapid and divergent evolution, and is likely to be a major driver of evolutionary change within species.

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Accession codes, primary accessions, embl/genbank/ddbj, data deposits.

The ancestral genome sequence of phage Φ2 has been submitted to the EMBL Nucleotide Sequence Database under accession number FN594518 .

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Acknowledgements

We are grateful to M. Begon and G. Hurst for comments on previous drafts of this manuscript. We acknowledge funding from the Natural Environment Research Council, the Wellcome Trust, the Leverhulme Trust and the European Research Council. We are grateful for technical assistance from staff at the Centre for Genomic Research, University of Liverpool.

Author Contributions M.A.B. and S.P. conceived and designed the study; T.V. performed selection experiments, infectivity assays and prepared samples for population genomic sequencing; B.L. and M.A.B. performed further assays; R.B. prepared samples for ancestral genome sequencing; N.R.T., M.Q., F.S. and D.W. performed ancestral genome sequencing and assembly; A.J.S. and N.R.T. performed genome annotation; S.P., T.V. and M.A.B. analysed the data; M.A.B., S.P., A.B., A.J.S. and N.R.T. wrote the manuscript; T.V. and N.H. commented on the manuscript; M.A.B., A.F. and S.P. supervised the research; S.P., N.H., A.B. and M.A.B. obtained funding.

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Steve Paterson, Tom Vogwill and Michael A. Brockhurst: These authors contributed equally to this work.

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School of Biological Sciences, Biosciences Building, University of Liverpool, Crown Street, Liverpool L69 7ZB, UK ,

Steve Paterson, Tom Vogwill, Ben Libberton, Andrew Fenton, Neil Hall & Michael A. Brockhurst

Zoology Department, University of Oxford, South Parks Road, Oxford OX1 3PS, UK,

Angus Buckling & Rebecca Benmayor

SIMBIOS Centre, Level 5 Kydd Building, University of Abertay Dundee, Bell Street, Dundee DD1 1HG, UK ,

Andrew J. Spiers

Pathogen Genomics, The Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, Cambridge CB10 1SA, UK ,

Nicholas R. Thomson, Mike Quail, Frances Smith & Danielle Walker

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Correspondence to Michael A. Brockhurst .

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Paterson, S., Vogwill, T., Buckling, A. et al. Antagonistic coevolution accelerates molecular evolution. Nature 464 , 275–278 (2010). https://doi.org/10.1038/nature08798

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Red Queen Hypothesis

The Red Queen Hypothesis, named after the Red Queen in the book Alice in Wonderland, brings together two evolutionary theories.

This article is a part of the guide:

• Law Of Segregation
• Darwin’s Finches
• Biology Experiments
• Transforming Principle
• Industrial Melanism

Browse Full Outline

• 1 Biology Experiments
• 2 Law Of Segregation
• 3 Darwin’s Finches
• 4 Industrial Melanism
• 5 Red Queen Hypothesis
• 6 Transforming Principle

The basis for the entire theory is down to ‘the evolutionary arms race’, where prey and predator constantly evolve together to reach some sort of uneasy balance.

An example of the Red Queen Hypothesis might be one of the plants that evolve toxins to kill off predators such as caterpillars .

If the plant, under predation selection pressure, evolved a new type of toxin to which the caterpillar had no immunity, most of the caterpillars would die off and the tree would flourish. This victory would be short lived, as only the caterpillars immune to the toxin, even if only a tiny percentage, would breed rapidly, and once again the tree would be under attack.

Sexual Reproduction and Genetics

For the Red Queen Hypothesis to happen, some sort of genetic mixing of genes must happen, such as sexual reproduction, as this throws up enough random fluctuations and mutations to allow new traits to appear. Most random mutations will disappear, as they confer no advantage or may even be detrimental.

Occasionally, they will give an advantage and will quickly spread through a population, as individuals with this genotype will have a competitive advantage. The industrial melanism shown by the peppered moth is an excellent example of this process in action. Without this mutation there would be a chance that the species may have become extinct.

If its population had shrunk, through predation or disease, to a small size, the species would have been open to environmental factors wiping it out.

Genetic fluctuations rely on probability and numbers. A large population is much more able, by chance, to throw up random mutations, whilst a small population is less likely.

Other Selection Pressures

Predator/prey relationships are not the only factors in the Red Queen Hypothesis.

If many species are competing for the same resources, mutations are sometimes needed to prevent a species from being out-competed. This is possibly one of the reasons why sexual reproduction occurs in higher species. If no random mixing occurred, then a bacteria or parasite may quickly evolve into a lethal form which would wipe out a species.

Sexual reproduction means that in a large population, there would be enough individuals with resistance to breed, pass the trait on and continue the species.

In a strange way, this benefits both host and parasite because, if a parasite or bacteria was so effective that it killed the host species, then it too is guaranteed extinction.

This process of sexual selection may explain why the vast majority of genes in vertebrates are dormant and do nothing (often called ‘junk DNA’) as they are preserving possible mutations that might suddenly be needed in the future if the environment or parasite pressure changes.

Sickle Cell Anemia

Often, these genes can even be detrimental. In humanity, sickle cell anemia is a gene that causes the red blood cells of an individual to become sickle shaped and less able to carry oxygen.

If both of an individual’s parents pass on the gene, the individual will have the full blown disease and, without medical intervention, probably die. Logically, it would be expected that this trait should die out of a population due to natural selection.

However, another factor has to be thrown in to the mix, malaria, where a parasite enters the blood stream through a mosquito bite. This parasite cannot live in blood affected by the sickle cell disease.

All of a sudden, the demographic changes; individuals with no copies of the gene are likely to suffer malaria, those with two copies anemia.

Only the individuals with one copy of the gene will be unaffected, as they do not have enough cells affected to cause the anemic effects, but enough to deter the malaria parasite, the cost being that some of their offspring will suffer anemia or malaria susceptibility.

In a final support to the Red Queen Hypothesis , this mutation has occurred independently, at least four times in human history, with the same gene involved, indicating that this gene must have been preserved for some reason and parasitic pressure caused it to be manifested.

Species, whilst improving and evolving to be more successful must face some sort of pressure from parasites or predators in order to evolve, with sexual reproduction being one of the main factors. Whole ecosystems and food chains are kept in check by this evolutionary ‘arms race’ as described by the Red Queen Hypothesis.

Sexual reproduction also acts as a safeguard against extinction. If a natural disaster or epidemic almost wipes out a species, a large population is genetically diverse enough to allow some individuals to survive and the species can once again prosper.

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Martyn Shuttleworth (Jan 1, 2008). Red Queen Hypothesis. Retrieved Sep 01, 2024 from Explorable.com: https://explorable.com/red-queen-hypothesis

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