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Edits to Wikipedia article Red Queen Hypothesis:

Further evidence was observed of the Red Queen to see allelic affects under sexual selection. The Red Queen Hypothesis leads to the understanding that allelic recombination is advantageous for populations that engage in aggressive biotic interactions, such as predator-host interactions. In cases of predator-host relations, sexual reproduction can quicken the production of new multi-locus genotypes allowing the host to escape invasion by of parasites that have adapted to the prior generations of typical parasites [15]. Mutational effects can be represented by mutational models to describe how recombination through sexual reproduction can be advantageous. According to the mutational deterministic hypothesis if the deleterious mutation rate is high, and if the deleterious mutations interact to cause a general decline in organismal fitness, then sexual reproduction provides an advantage over asexually reproducing organisms by allowing populations to eliminate the deleterious mutations not only more rapidly, but also most effectively [16]. Recombination is one of the fundamental means that explain the reason many organisms have evolved to reproduce sexually.

The name of the Red Queen Hypothesis was given its name by Lewis Carroll in The Looking Glass. Van Valen coined the hypothesis “Red Queen” because under this interpretation where species have to “run” or evolve in order to stay in the same place or remain extant. The phrase “Red Queen Hypothesis” comes from the second chapter of Carroll’s novel Through the Looking Glass. In the main character’s dream, Alice, finds that objects appear left-to-right, as if in a mirror. Alice then notices the chess pieces are alive, most notably the Red Queen. The Red Queen explains that she is in a world different that her own. At the top of a hill, the Red queen begins to run, and Alice begins to chase after her. Alice is confused by the fact that even though they are running, they are staying in the exact same place. Alice asked the queen why this is and the Red Queen remarks, “Now here, you see, it takes all the running you can do to keep in the same place[5]. This is comparable to the evolutionary arms race seen in nature where organisms need to stay constantly evolving to not “lose” the race with their predators or parasites.

FINAL PAPER The Red Queen Hypothesis

The Red Queen hypothesis is an evolutionary principle that describes the phenomenon that organisms are evolving at the same rate as their predators. These organisms are in a constant co-evolutionary arms race with their predators, parasites, and pathogens. Organisms are going to want to evolve at a faster rate to evade the negative selective pressures that their biological enemies are imposing. These enemies on the other hand are evolving as fast as their prey to catch up with the organisms (Lively). These two different organisms are in a race resulting in evolution at the same time as the other. This concept is critical in understanding predator-prey relationships and also parasitic relationships. Van Valen coined the hypothesis “Red Queen” because under this interpretation where species have to “run” or evolve in order to stay in the same place or remain extant (Lively). The phrase “Red Queen Hypothesis” comes from the second chapter of Carroll’s novel Through the Looking Glass (Carrol 1872). In the main character’s dream, Alice, finds that objects appear left-to-right, as if in a mirror. Alice then notices the chess pieces are alive, most notably the Red Queen. The Red Queen explains that she is in a world different that her own. At the top of a hill, the Red queen begins to run, and Alice begins to chase after her. Alice is confused by the fact that even though they are running, they are staying in the exact same place (Carrol 1872). Alice asked the queen why this is and the Red Queen remarks, “Now here, you see, it takes all the running you can do to keep in the same place (Carrol 1872). This is comparable to the evolutionary arms race seen in nature where organisms need to stay constantly evolving to not “lose” the race with their predators or parasites. The Red Queen Hypothesis is used to describe two similar ideas of co-evolution resulting in an increased probability of extinction and oscillation of genetic frequencies over time between organisms. The original evolutionary thought is that coevolution could lead to scenarios in which the chance of extinction is somewhat constant over millions of years (Van Valen 1973). In coevolving interactions the evolutionary change could happen in either the prey or the host and the likelihood that those changes are not dependent of the species age (Van Valen 1973). The other idea of the Red Queen Hypothesis is that during the organisms co-evolving, there is a chance that organisms have sustained oscillations in their genotypic frequencies. This idea forms the basis of one of the leading cases of why sexual reproduction is advantageous. Within species that reproduce asexually, co-evolutionary interactions with parasites may in turn select for sexual reproduction to lower the risk of infection of the next generation (Lively). There have been many important implications on how the Red Queen Hypothesis can be applied to explain the evolution of sex. Many plant and animal species reproduce sexually, but there are many costs associated with this form of reproduction. Genetic recombination might out weigh the costs of sexual reproduction since recombination helps elevate pressure of predators by creating new genotypes (Salathé, et al 2009). There are computer simulations to measure mathematical models to measure host-parasite coevolution. Through research on the Red Queen hypothesis it is found that the real driving force underlying the evolutionary arms race is neither the immediate organism, nor the offspring of the organism, but a delayed short-term effect (Salathé, et al 2009). Understanding the Red Queen hypothesis can lead to understanding the benefits of sexual reproduction. Mutational and ecological effects can aid in explaining the advantages of sex. It is crucial to understand the selective forces that drive sexual reproduction (Salathé, et al 2009). The Red Queen Hypothesis leads to the understanding that allelic recombination is advantageous for populations that engage in aggressive biotic interactions, such as predator-host interactions. In cases of predator-host relations, sexual reproduction can quicken the production of new multi-locus genotypes allowing the host to escape invasion by of parasites that have adapted to the prior generations of typical parasites (Cooper, et al 2005). Mutational effects can be represented by mutational models to describe how recombination through sexual reproduction can be advantageous. According to the mutational deterministic hypothesis if the deleterious mutation rate is high, and if the deleterious mutations interact to cause a general decline in organismal fitness, then sexual reproduction provides an advantage over asexually reproducing organisms by allowing populations to eliminate the deleterious mutations not only more rapidly, but also most effectively (Cooper, et al 2005). Recombination is one of the fundamental means that explain the reason many organisms have evolved to reproduce sexually. One of the best ecological models to explain sexual reproduction being an advantageous mode of reproduction can be described by the geographic mosaic theory. The geographic mosaic theory of co-evolution states that populations of interacting species can produce selection mosaics established as co-evolutionary “hot spots” and “cold spots.” (King, et al 2009). The frequency of sexual reproduction of freshwater snails was observed by the presence of hot spots and cold spots with coevolving parasites. The most common places for sexual reproduction of snails, shallow-water areas of lakes, were deemed hot spots, while the deeper habitats are cold spots. The results supported the geographic mosaic theory in which the intensity of selection results drastically from location to location (King, et al 2009). The results from the snail reproduction study supports the Red Queen hypothesis in that sex is associated with co-evolutionary hot spot spots for active parasites. This is shown by the co-evolution of snails and its parasite. The Red Queen hypothesis explains the evolutionary advantages of sexual reproduction through mutational and ecological models. The Red Queen hypothesis can be seen all throughout nature. There are examples of cells within organisms, eukaryotic and prokaryotic creatures, and even RNA viruses that are participating in the evolutionary arms race with parasitic organisms or prey. One example of the Red Queen hypothesis seen within the environment is lymphocytes in our body. Development of T-cells has been altered over time. The survival of these cells has been described as an active process, dependent on the expression of particular sets of resting T-cells. Research has identified Kruppel-like zinc-finger transcription factor called lung Kruppel-like factor (LKLF) that is expressed in mature T-cells but not in immature lymphocytes (Freitas, et al 1997). The presence of the transcription factor LKLF is vital for the survival of naīve T-cells. These are the cells that are responsible for responsible against foreign antigens. Freitas, et al observed that the Red Queen Hypothesis was observed suggesting that the peripheral immune system, ran by these naīve T cells, is “running” against pathogens (1997). Along with lymphocytes being seen to exhibit the Red Queen, Daphnia retrocurva and RNA viruses have also been seen to add claim to this hypothesis. A study observing the prey portion of the Red Queen hypothesis was performed retrieving diapausing eggs from lake sediments to observe the evolutionary response through environmental changes (Kerfoot, et al 2004). The predator Daphnia retrocurva was also observed to study its evolution with its prey, the diaspausing eggs. The results confirmed the Red Queen hypothesis in which the egg hatchlings documented micro evolutionary adjustments supporting continual evolution in prey against differential balance of predication (Kerfoot, et al 2004). The Red Queen hypothesis is also seen within RNA viruses. An experiment by Clarke, et al studied two clonal populations of vesicular stomatitis virus of equal relative fitness that were mixed and allowed to compete in transfers in vitro as large virus populations. Over time, one or the other population excluded its competitor population, even though both the winners and losers exhibited gains in fitness (Clarke, et al 1994). This study of viruses exhibits the Red Queen Hypothesis in how two organisms are constantly co-evolving with each other. Mathematical and statistical models are important in observing how the Red Queen hypothesis acts over time and how it affects the organisms. This hypothesis addresses when the homogenous taxonomic group is among the same species. Recent approaches to solving the problem of the varying statistical approaches against taxon age and rate of extinction are being combatted by Weibull and Exponential modeling, and Cox’s proportional hazards modeling (Wiltshire, et al 2014). Wiltshire, et al observed Planktonic Foraminifera extinction data to compare the taxonomic age with the rate of extinction this organism went through (2014). By designing new statistical means of interpreting data allows for greater understanding behind the mechanisms of the Red Queen hypothesis. There are many examples of the Red Queen hypothesis being displayed within nature, but there are scientists that argue that this hypothesis makes certain assumptions that are false. The Red Queen hypothesis’s basis is that: evolutionary adaptations are continuous, organisms are important agents of extinction, and evolution is a zero-sum process (Vermeij 2013). It is debated that even though mutations are considered frequent enough to be continuous, adaptations cannot be empirically stated as being continuous. Van Valen stated the longer the taxon’s age the more likely that organism was to become extinct but this was more recently disproven. And lastly there are multiple factors that go into an organism evolving (Vermeij 2013). These are interesting points stated that are encouraging scientists to continue the research into the Red Queen hypothesis and its implications on the environment. The Red Queen hypothesis is a theory that states that organisms are in a constant evolutionary race against parasites and prey that leads them nowhere. Ecological and mutational models have been shown that sexual reproduction is more advantageous than asexual reproduction due to allelic recombination, becoming an important result of this hypothesis. The Red Queen is seen in a diverse set of organisms that lead them to remain extant. The study of sex and the Red Queen hypothesis is one that is a continual area of research with more evidence brought to light each year.

Three suggestions to the Red Queen Hypothesis Wikipedia page. 1.) In this article I feel that adding graphics of data graphs or pictures of species that demonstrate examples of the Red Queen Hypothesis would add to the reader's comprehension of this evolutionary biological hypothesis. Mckinney.264 (talk) 04:26, 1 October 2014 (UTC) 2.) There has been countless research completed by evolutionary biologists in this field of evolutionary biology. Would it be possible for in the Historical Studies section that there be more research explained on studies looking at the results? Mckinney.264 (talk) 04:22, 1 October 2014 (UTC) 3.)I agree, I think that there needs to be more of a context of where the name Red Queen Hypothesis is derived from. I know within the page it discusses Lewis Carrol's origin,but maybe a sentence or more will help bring initial context to the piece. Mckinney.264 (talk) 04:14, 1 October 2014 (UTC)

Url with added sentence: https://en.wikipedia.org/wiki/Red_Queen_hypothesis

Sentence added with citation: Van Valen's metaphor can be explained by as one species increases their fitness becoming better adapted to their environment, other species in which that species interacts with will become adversely affected.[3] Vermeij, Gerrat; Roopnarine, Peter (2013). "Reining in the Red Queen: the dynamics of adaptation and extinction reexamined". Paleobiology 39 (4): 560–575.

Topic: Red Queen Hypothesis

Stephen McKinney Recitation: Thursday 9:10 Annotated Bibliography

Clay, K., & Kover, P. X. (January 01, 1996). The Red Queen Hypothesis and plant/pathogen interactions. Annual Review of Phytopathology, 34, 29-50.

Red Queen Hypothesis examines the path that pathogens take to maintain sexual reproduction within hosts. This is based on the assumption that parasites become specialized on common host genotypes. This will in turn reduce the fitness of the affected host. The specific conditions in which the hypothesis thrives in were observed to lower the fitness of the host the greatest.

Freitas, A. A., & Rocha, B. (September 26, 1997). Lymphocyte survival: a red queen hypothesis. Science (New York, NY), 277, 5334.)

This article by Freitas looks at lymphocyte’s life cycle and observes the evolution of how it has progressed in eukaryotic organisms. It examines how the development of T-cells has been altered over time. There were experiments ran to observe specific interactions with the cells. With understanding the functions it will be able to understand the evolution of the cells, thus supporting the Red Queen Hypothesis.

Salathé, M., Kouyos, R. D., & Bonhoeffer, S. (July 01, 2009). On the causes of selection for recombination underlying the red queen hypothesis. The American Naturalist, 174, 31-42.

In this article it discusses how through sexual reproduction there is great cost, but genetic recombination might outweigh these costs. Short or long term effects can drive the selection. Computer simulations can model the coevolution of host-parasite co-evolution to measure a variety of factors. The results show the driving force of the Red Queen Hypothesis is delayed short-term effects. This paper is critical in determining the force that supports the Red Queen Hypothesis.

Vermeij, G. J., & Roopnarine, P. D. (January 01, 2013). Reining in the Red Queen: the dynamics of adaptation and extinction reexamined. Paleobiology, 39, 4, 560-575.

This article examines three false assumptions made by this hypothesis. They are that evolutionary adaptation is continuous, organisms are important agents of extinction, and evolution is zero-sum process in which living things divide up unchanging quantity of resources. It shows that the Red Queen Hypothesis reigns in only few circumstances.

Wiltshire, J., Wiltshire, J., Huffer, F. W., & Parker, W. C. (April 01, 2014). A general class of test statistics for Van Valen's Red Queen hypothesis. Journal of Applied Statistics, 41, 9, 2028-2043.

This is a mathematical approach to understanding the Red Queen Hypothesis. Wiltshire and colleagues uses a mathematical approach to testing and further explaining Van Valen’s hypothesis. The taxon age and rate of extinction are the two primary variables being observed. By understanding a statistical approach it will better validate the conceptual points of the hypothesis.