Evolution and Natural Selection I have called this principle, by which each slight variation, if useful, is preserved, by the term Natural Selection - Charles Darwin, The Origin of Species Evolution and Natural Selection We wish to ask: • What are the main points of Darwin’s theory of evolution? • How did observations of nature lead to the formulation of the theory of evolution? • How does the process of natural selection work? • What evidence do we have for local adaptation? • How can natural selection affect the frequency of traits over successive generations? The Intellectual Climate in Darwin’s Time • The age of the earth was uncertain, but studies of strata suggested a time sequence. • Uniformitarianism -- the concept that present conditions and processes are the key to the past -was becoming influential. • Fossil discoveries revealed unknown life forms, suggesting extinctions had occurred. • Similarities among organisms were beginning to be seen as evidence of relatedness. Jean Baptiste Lamarck • A predecessor of Darwin, Lamarck argued that animals adapted to their environment • His advocacy of gradual evolution change is remembered mainly because he got the mechanism wrong. Darwin’s Theory of Evolution • Organisms have changed over time. Those living today are different from those that lived in the past. • All organisms are derived from common ancestors by a process of branching. Similarities of traits are evidence of a recent, common ancestor. • Change is gradual and slow, requiring a very long time. • Natural selection is the mechanism of evolutionary change. The Process of Natural Selection • All populations have the capacity for geometric increase, yet do not. • There must be a “struggle” to survive and reproduce, in which only a few are successful. • Organisms vary in traits that influence their likelihood of success in this “struggle”. • Organisms whose traits enable them to survive and reproduce will contribute a greater number of offspring to the next generation. • Offspring resemble their parents, including in these traits that influence survival. • Therefore the population in the next generation will include a greater fraction of individuals with whatever traits enabled their parents to survive and reproduce. Natural selection requires….. For natural selection to occur, two requirements are essential: 1. There must be inherited variation for some trait. Examples: beak size, color pattern, fleetness. 2. There must be a greater or lesser probability of survival and reproduction due to possession of that trait. Examples: peppered moth, Galapagos finches. The peppered moth, Biston betularia. Light and dark morphs on (a) a light background, (b) a dark background Distribution of the two moth morphotypes during the industrial era The Link between Evolution and Genetics “Evolution is the change in the genetic composition of a population over time” • Where does genetic variation come from? - Mutation (raw material, ultimate source) - sex, meiosis, and exchange of chromosome segments - other chromosomal re-arrangements • What are other causes of changes in the genetic composition of a population over time? – Non random mating - migration – Mutation - natural selection – “Sampling error or genetic drift” The Hardy-Weinberg Equilibrium “In a large population, with random mating, and in the absence of forces that would change their proportions, the proportions of alleles at a given locust will remain constant” Allele: Frequency: A p a q The H-W Equilibrium is significant because: 1. It tells us that as long as mating is random and nothing else happens, gene frequencies will stay unchanged indefinitely 2. It gives us a tool to calculate allele frequencies if we know genotype frequencies, and vice-versa Three Types of Selection Envision a frequency distribution for a trait such as beak size, much like a grade distribution. • Under stabilizing selection, extreme varieties from both “tails” of the frequency distribution are selected against; the mean stays constant. • Under directional selection, individuals at one tail of the frequency distribution are favored (“selected for”), the other tail is selected against. The mean value shifts in the direction favored • Under diversifying (disruptive) selection, both extremes are favored at the expense of intermediate varieties. The result is a bimodal distribution. • Stabilizing directional diversifying • Birth weight appears to be under stabilizing selection. Babies that are too large or too small have higher mortality • Directional selection in horse evolution. Over time, the horse grew taller and its hoof evolved as the nail of the middle digit. Its molars grew larger as the horse became a bigger animal, and its diet became primarily grasses. A drier climate resulted in changes in habitat (open savannahs instead of forests) and food (grass instead of leafy plants). Darwin’s finches evolved from a common ancestor, similar to a species now found on the mainland of Ecuador. We’ll look at this adaptive radiation again when we discuss how species compete for food. Sub-species of the rat snake Elaphe obsoleta, referred to as geographical “races”. They interbreed where their ranges meet, so are not distinct species. A phylogenetic tree constructed from similarities between cytochrome c molecules in various organisms. Summary • The core of Darwin’s theory of evolution is its mechanism -- the theory of natural selection. • Natural selection requires inherited variation in a trait, and differential survival and reproduction associated with possession of that trait. • Examples of natural selection are well documented, both by observation and from the fossil record. • Selection acts on the frequency distribution of traits, and can take the form of stabilizing, directional, or diversifying selection.