Jenna Peon BSC 1005 Evolution Term Paper INTRODUCTION George R.

Price lived from 1922 to 1975, carrying out most of his life in the United States before moving to London in 1967. Though his contributions to evolution and science in many other areas were extensive and astonishing, he has managed to remain a fairly obscure figure in history. The typical person has never heard of Price, nor his elegant equation that changed the way that modern evolutionary biology is studied. He was educated as a chemist and even worked on the Manhattan Project studying uranium enrichment; in display of his versatility he also briefly tried out the field of journalism. Among his other professional projects include relationships with companies like IBM and involvement in developing cancer radiation technology (Schwartz, 2000). He wandered into the field of evolutionary biology later on in his career, simultaneously going through extreme changes in his personal and religious life. Though Price was never formally educated in the field of biology, he made several lasting and prominent contributions to the field. Among these accomplishments is the creation of the legendary Price Equation, which is a mathematical theorem that successfully captures the evolutionary processes and explains how all traits, whether physical or mental, may be passed on through genetics and evolve over time. He formulated this sophisticated yet incredibly simple theorem after reading an article by Professor W.D. Hamilton about altruism and its genetics. This discovery would change the rest of Prices life (Rahul, 2012). Another thing that Price did was perfect R.A. Fishers fundamental theory of natural selection in a way so that people could actually understand and apply it (Frank, 2002).

The most relevant of Prices major contributions to the field of evolution is his allencompassing Price Equation. It was able to capture the evolutionary processes, both the deterministic and the random, and has changed the way that evolutionary biology is understood and studied. The equation reads as follows in its most common form: (
)

Equation 1

The equation itself represents the change due to differential survival and reproduction plus the change resulting from processes involved in reproduction (BSC 1005 notes, 10/10/2013). RESULTS Phenotype, which is represented by the Greek phi, is the specific trait that is being studied. Fitness, depicted by the letter omega () is essentially just the number of offspring produced. The first component in Prices theorem, the covariance between phenotype and fitness, represents the change due to selection (deterministic, non-random). The second component of Equation 1 represents the change due to transmission (stochastic, random). In this paper, only the non-stochastic processes will be focused on. Because evolution by natural selection assumes that there are perfectly heritable (=0) phenotypes and an expected fitness value, we will then dismiss the second component of Prices Equation by setting it to 0, and be able to focus on only the change due to selection. The lowercase delta is defined as: ,

which is the average change between the parent and child phenotypes. The omega, or relative fitness, () is defined as , where is fitness and is the per capita growth rate for the parent population. So here, delta omega bar in the Price Equation (the product of the average of delta

and the average of omega) are essentially cut out in this study due to the focus on only the deterministic processes (BSC 1005 notes, 10/24/2013 and 10/29/2013). In Equation 1, phi bar represents the mean offspring phenotype of the parent. The upper case delta represents change. Thus we have delta phi bar ( ) meaning the change

in the average phenotype. In evolution, this change is measured from one generation to the nextmeaning from the parent phenotype to the respective offspring phenotype- which are denoted as and respectively. Delta phi bar can also be expressed as an identity that is defined as the

difference in the average phenotype of the offspring and the average phenotype in the current (or parent) generation, denoted as follows: Equation 2

The first term in Equation 2, phi bar prime (average phenotype in the next generation), is calculated by dividing by the sum of the fitness from the parents (). The second term, phi

bar (average phenotype in the current generation), is calculated by dividing the sum of the parent phenotypes ( 10/03/2013). Within itself, this principle of his theorem proves that there must be reproduction in order to capture evolution, because without reproduction (=0), phi prime would be equal to , which by the laws of mathematics would make evolution undefined. If there is 0 fitness, meaning no production of offspring, then there is no evolution. This leads to the conclusion that the linear slope of the relationship between fitness and phenotype must be nonzero (BSC 1005 Notes, 09/03/2013). by the size of the population of which the parent is a part (n) (BSC 1005 notes,

A covariance is a function that is used when you are taking the average of the product of two values; it is a measure of how one variable will change as you change the other, or essentially a measure of the variables dependence upon one another. Price defined evolution by these covariance terms in his equation. It is the sole most important function in Prices theorem and is an astonishingly simple linear regression relationship that was able to explain many of the questions of earlier evolutionary scientists. Covariance captures the directional behavior of a relationship between two differing values, and therefore is not useful in numerical calculation that represent actual figures related to the relationship, but is more correlative in nature (Price, 1970). The slope of this line that is mentioned in the above paragraph can be defined as: Equation 3

(BSC 1005 Notes, 10/15/2013). A covariance value can be positive or negative. This will determine whether the relationship between the two terms is positive or negative. In a positive relationship, one term will increase as the other is increased or likewise decrease as the other is decreased. In a negative covariance relationship, one term will increase as the other is decreased, or vice versa. Positive slope allows for the implication that the population is evolving towards higher phenotypes, and negative slope shows just the opposite (BSC 1005 notes, 10/29/2013). In order to estimate these covariance terms from a natural population, there would have to be extensive study on multiple sample populations in laboratory settings to estimate, or the study could be done directly in the field, testing sample subjects directly from the natural population.

DISCUSSION Adaptations are the result of the causality that is implied by this covariance association between phenotype and fitness that we see in Equation 1. So essentially, adaptations occur when natural selection is acting in a population; and likewise, there is always going to be a causal factor that is leading the population towards the specific trait or phenotype that is displayed or studied. Over many generations, the organisms which have these adaptive traits (and therefore are undergoing the process of evolution) begin to make up more and more of the population (BSC 1005 notes, 09/03/2013). This means two things: first, that evolutionary processes are at work within the population being studied and are altering the genetics from the current generation to the next, and also that the fitness of individuals carrying the adaptive trait is probably increasing over time. Applying the Price Equation to a natural population can be done in just about any way, as long as there is a population which reproduces. The application must be carried out in a precise method, substituting values for each of the covariance terms that are proportional to their relationship. The most important part of a study of this manner is correctly defining each individual term, so as not to upset the equation as a whole nor the calculation of your final covariance term. A study of evolution within any population must be carried out in a way so that only the desired evolutionary processes are zeroed in on. In the case of this analysis, only the deterministic processes, like natural selection, are being considered. The environment and other evolutionary processes like drift and mutation must be accounted for and removed from the equation. There must be careful attention paid to the exact values chosen to represent the fitness

and other components, as they are all relative to one another in a linear regression equation such as this one. Personally, I would be interested in studying an example that has a personal importance to me, much like George R. Price studied anything that sparked his interest. Being that I was diagnosed with Type One Diabetes almost 6 years ago, I have always had a profound curiosity about how prevalent it is, what causes it, and how my carrying of this gene may or may not affect my offspring and even my distant relatives in the future. The advanced scientific nature of our contemporary society has begun to delve into these questions, but have not even touched the surface due to the fact that only about 60% of the genetic causal factor has been discovered (Skyler and Sosenko, 2013). There is obviously a genetic factor (causal factor, essentially an adaptation/mutation) that links the possibility of Type One Diabetes occurring to specific genes or allele frequencies, but these have yet to be identified. If the origin of this autoimmune disease could be narrowed down, the Price Equation can be used, as it is in the 1972 article Extension of covariance selection mathematics to measure allele frequencies as a covariance term (Price, 1972.) It would be quite interesting to me to work on researching this specific problem, mostly because it is such a close personal issue to my life. Also, it intrigues me because it is something that scientists have not been able to figure out yet despite the huge technological advances and availability of funds for research as this disease continues to affect thousands more each year. Though further research is needed for the Price Equation to be able to be applied here, that would be the ideal example that I would choose to study if I had the chance, as it could be used to help potentially find a way to delay or stop the onset of juvenile diabetes.

Using the Price Equation to estimate allele frequencies can be done by using different derivations of the equation, as seen in the article by Marco Antonio on theoretical evolutionary genetics. It can be done through a few different methods, but you can use the Price Equation method for deriving the change in allele frequency over time (Antonio, 2013). I would definitely be interested in doing a longitudinal study of subjects who have signs of developing diabetes or a genetic predisposition to the disease, to see how many are actually affected by it. In turn, I would like to study their offspring, to measure how the allele frequency changes over time and thus be able to hypothesize how my own offspring and relatives might be affected, and how the genetics of the disease themselves can or will evolve over time. The Price Equation can be and is being applied in so many different ways, many of which I am sure George Price could not even imagine in his day. The scientific world is expanding exponentially with each passing day, yet we are still building upon the solid foundations which great scientists like Darwin, Fisher, Hamilton, and Price have laid down for us in history.

REFERENCES Frank, S. A. (2002). George price. In M. Pagel (Ed.), Encyclopedia of Evolution (pp. 930-931). Oxford University Press. Retrieved from http://stevefrank.org/reprints-pdf/02Price.pdf Papadopoulos, A. Notes taken in class from 08/2013 to 12/2013. BSC 1005-1196 Price, G.R. (1970) Selection and covariance. Nature 227:520-521. Retrieved from http://www0.cs.ucl.ac.uk/staff/wlangdon/ftp/papers/price_nature.pdf Price, G.R. (1972) Extension of covariance selection mathematics. Annals of Human Genetics 35:485-590.

Rahul K. (2012, October 15). Killed by Kindness [Video file]. Retrieved from http://www.youtube.com/watch?v=WYdgcKjbp3Y Schwartz, J. (2000). Death of an altruist. Lingua Franca, 10 (5), Retrieved from http://download.bioon.com.cn/upload/month_0807/20080730_b5faf5d6b599cdd40d7019 R7kxgaLm5c.attach.pdf Skyler, J., & Sosenko, J. (2013). The evolution of type 1 diabetes. Journal of the American Medical Association, 309(23), 2491-2492. Retrieved from http://jama.jamanetwork.com/article.aspx?articleid=1697946