BIO 140 3rd exam!!

1920s genetic researches on mutations Resulted to an alternative to Darwins Gradual evolution Evolution occurred in rapid leaps as a result of a radical phenotypic changes caused by mutations Saltatory evolution Gaps in the evolution in higher taxa. Saltation evolutionary jumps. Gaps between taxa. Remaining gaps in Darwins concept of hereditary variation 1. Mutation alone could not fully account for the observed hereditary variation. 2. Mendels laws of inheritance did not account for the characters that vary along a continuum in a population (continuous variation) Qualitative traits: Discrete/distinct variation Population Genetics Turning point in evolutionary ideas (1930s) It showed the extensive genetic variation within population and the evolutionary factors that affect it. It recognized the importance of quantitative inheritance (quantitative genetics) It reconciled Darwinism and Mendelism Darwinism Modified to Modern Evolutionary Synthesis Theory Contributors: 1. Theodosiu Dobzhansky (geneticist) 2. Ernst Mayr (biogeographer and taxonomist) 3. George C. Simpson (paleontologist) 4. G. Ledyard Stebbins (Botanists) Modern Synthesis Theory 1. It integrated Mendels principle of inheritance as source of hereditary variation Polygenic quantitative traits 2. It emphasized the importance of population as the units of evolution

Characteristic Life Span

Population Immortal (presence of perpetuation) Genetic makeDoes not change Can change from up generation to generation 3. It defined the role of natural selection as an important mechanism of evolution 4. It described how gradualism can account fro large changes through accumulation of small change over long periods of time. WHAT IS POPULATION??? Population is a localized groups of organism which belong to the same species Species groups of actually of potentially interbreeding individual which are reproductively isolated from other such groups Gene Pool The total aggregate of genes in a population at any one time Consists of all the allele at all gene loci in all individuals of a population Alleles from this pool will be the source of the offsprings of the next generation For diploid individuals Each locus has 2 copies or allele of a gene, e.g. A and a (assume A is dominant) These alleles combine to form a genotype An allele is said to be fixed in the gene pool if all members of the population are homozygous for that allele E.g. frequency of A ~f(A) = 100% or 1 Genetic Structure of a Population Refers to a populations frequencies of allele and genotype: 1. Allele Frequency the relative proportion of an allele (in a given locus) in a population 2. Genotypic frequency the proportion of individuals bearing a given genotype (at a locus) Letter Symbols for Frequencies Gene Frequencies F(A) = p F(a) = q Genotypic Frequencies (at equilibrium) F(AA) = p2 F(Aa) = 2pq F(aa) = q2

Individuals Mortal

Ex: Given a population of wildflowers where in 20 plants have white flowers 160 have pink flowers and 320 have red flowers. Calculate the gene and genotypic frequencies (assume the population is in equilibrium. There is incomplete dominance) Trait: Flower color (Let A = allele for red and a = allele for white) [Type a quote from the document or the summary of an interesting point. You can position the text box anywhere in the document. Use the Text Box Tools tab to change the formatting of the pull quote text box.]

MICROEVOLUTION???? Evolution that occur at or below the level of species Such as a change in the gene frequencies of a population of organisms. Also the process by which new species are created (speciation) Maybe due to reversal processes: mutation, gene flow, genetic drift and natural selection. Examples of Microevolution Bacterial strain that have become resistant to antibiotics Color change in moths over time Because microevolution can be observed directly, it is widely accepted (unlike microevolution ~ garnered controversies) In the absence of other factors, the segregation and recombination of allele during meiosis and fertilization will not alter the genetic make-up of a population. HARDY and WEINBERG LAW (1908) Hardy (math professor at Cambridge) Weinberg (German physician) Hardy-Weinberg equilibrium In a large, randomly mating population, the allele frequencies p and q (For allele A and a) will attain in one generation the frequencies p2, 2pq and q2 for the three genotypes (AA, Aa, and aa), regardless of the initial genetic constitution and will remain in these proportion in the absence of selection, mutation and migration. When the five conditions in a population is met, then the population is stable, not evolving. Population genetics is the branch of biology, that provides the mathematical structure for the study of the process of microevolution.

Gene Frequencies: 3 methods 1. Assume Equilibrium

2. By basing it on the genotype frequencies

3. By allele counting

Hardy-Weinberg Model Describes the genetics of an ideal, NON evolving population This situation (rarely) exists in nature In actual populations, several factors can upset the Hardy-Weinberg equilibrium and cause evolutionary change ~ Evolutionary change

The larger the population, the less important is the effect of genetic drift. Even though natural population are not infinitely large, mostly are large enough that the effect of genetic drift is negligible In population with less than 100 individuals genetic drift can play a role in microevolution

Microevolution can be cause by:

1. 2. 3. 4. 5. Genetic Drift Migration or Gene flow Mutation Natural selection Non-random mating Leads to accumulation of favorable adaptations in a population The others are non-adaptive (and are called) non Darwinian changes.

2 situations which result in decrease in population size and many may cause genetic drift: 1. Bottleneck effect 2. Founder effect Bottleneck effect Genetic drift which results from drastic reduction in population size Due to natural disasters (by chance. Some individuals survive) The small surviving population is unlikely to represent the genetic variability in a population since some allele may be: 1. Under represented 2. Over represented 3. Totally absent For Example, a population of Northern Elephant seals as reduced to 20 individuals by hunters in the 1890s. They have been protected and the population has increased to about 50,000 Researchers found NO VARIATION exists in all the 24 gene loci examined in the present population A simple allele HAS BEEN FIXED at each of the 24 gene loci to genetic drift by the bottleneck effect. An allele is fixed in the population when its frequency is equal to 1.0. AFLP amplified fragment length polymorphism No variation in DNA bands after gel electrophoresis Polymorphic Locus many forms=many alleles Founder Effect Occur when a few individuals colonize a new habitat ~ founders Founder effect is genetic drift in a new colony The smaller the founding population the less likely its gene pool will be representative of the original populations genetic make-up Examples: Amish people in Pennsylvania USA (inbreeding, Ellis van Crevald syndrome) 1. Evolutionary divergence of the Galapagos finches

1. GENETIC DRIFT
Change in the gene pool of a small population due to change I.e. Chance events may cause the frequencies of allele to drift randomly from generation to generation.

Sampling error can happen The existing gene pool may not be accurately represented in the next generation.

Actual Examples 1. Change in gene frequencies of beetles 2. Change I cypress seed shape 3. Evolution of the white footed mouse The total population size has an effect on genetic drift Low/decrease in population size = High/increase effect on population High/increase population size = Low/decrease effect on population

2. High frequency of retinitis pigmentosa in human population of Tristan da Cunhal (Atlantic islands) 3. High deaf population in Marthas vineyard Inbreeding results in the expression of recessive genes

Examples: A wind storm may blow pollen from one plant population to another The migration of humans to different populations Effect: gene flow tends to reduce differences between population

2. Gene flow or Migration

The movement of fertile individuals between populations The transfer of gametes (and alleles they carry) between population Immigration and emigration Can change the frequencies of the native populations defending on: a) Number (frequency) of migrants~ m b) Number of generation of migration ~ n c) Difference in gene frequencies of between population The resulting q in the native population after n generations of migration (qm) can be computed.

3. Mutation
Change in the genetic material Can be: a) Point mutation (base mutation, frameshift) b) Change in chromosome structure (deletion, duplication, inversion, translocation) c) Change in chromosome number (aneuploidy, euploidy) The original source of genetic variation which is the new material for evolution Mutation tends to balance the effect of natural selection by generating new alleles In gametes, changes the gene pool of a population (especially one way or unidirectional mutation) Forward mutation = Aa Backward mutation = a A But has little quantitative effect on large population in a single generation. Neutral Theory of Molecular Evolution States that the majority of evolutionary change at the molecular level is caused by random drift as selectively neutral mutants Not all genetic variation is adaptive Variant alleles at a locus may confer no selective advantage or disadvantage Mutation rates in Gametes One mutation per 105 to 106 gametes Vary depending on the locus and species Some occur bi-directionally with different rates for forward (u) and backward (v) mutation

Where: m = frequency of migrants n = number of generations of migration qo = frequency of per allele in the native population Q = frequency of the allele in the migrant population

4. Non-Random mating
Increase the number of homozygous loci in population 2 types: a) Inbreeding (will not change frequency but increases homozygous genes) b) Assortative mating (increases gene frequency)

a. Inbreeding Individuals usually mate with close neighbors rather than with distant members of a population - occurs when population members do not dispose widely. - Mating between relatives or individuals bearing the genes identical by descent from a common ancestor b. Assortative mating select the mate - Individuals mate with partners like themselves in certain phenotypic characters - Like with like -may also increase homozygosity (AAxAA, AAxAa, AaxAa) Self-fertilization most extreme example of inbreeding Inbreeding results in genotypic frequencies (p2, 2pq and q2) that deviate from the frequencies predicted for Hardy-Weinberg equilibrium. One effect: frequency of homozygous recessive phenotypic increases

How genetic variation is preserved?? 1. Diploidy - Balanced polymorphism Favoring heterozygotes Heterozygote advantage heterozygote have greater reproductive success over homozygous individual Hybrid Vigor heterozygotes exhibit phenotypes that exceed that of the homozygous parent. Adaptive evolution is garnered by: A. Fitness a) Relative fitness b) Selection coefficient B. Modes of natural selection a) Stabilizing b) Directional c) Diversifying or disruptive Relative fitness (w) Reproductive coefficient Selection coefficient (s) Represents degree of selection When s=0 then w=1 Thus w=1-s Example: there is a 20% selection on the recessive individual in the population Fitness AA Aa aa s 0 0 0.2 w 1 1 0.8 Selection Against Extremes a) Stabilizing selection (favors heterozygotes)

5. Natural Selection
Differential success in the reproduction of different phenotypes Results from the interaction between organisms (of a population) with their environment Can change relative allele frequencies in the population evolution Some variants in a population leave more offsprings than others The HW equilibrium condition that all individuals in a population have equal ability to reproduce viable offspring is never met. Example: Biston betularia (peppered moth) Exhibits industrial melanism Classic example of natural selection (change in environment) Observed in Britain: Before 1850s: white After 1850s: Black Lichen covered Trunk Non-polluted areas Lichen-covered trunk Dark moths seen and eaten White moths flourish Lichen-free, soot covered trunk Industrial areas Soot covered trunk White moths seen and eaten Dark moths flourish

Example: Sickle Cell Anemia Let H: normal Hb (normal RBC) H: sickle Hb (RBC sickle)

b) Diversifying/ disruptional: Selection against the mean population

Morphospecies Species defined by their anatomical features Used by Linnaeus Still the most common method used today Difficulties: 1. It is difficult to determine if a set of organism represents multiple species or a single species with extensive phenotype variation 2. Two population morphologically similar are almost indistinguishable MAY BE different species based on other criteria Biological Species (Mayr, 1942) Population on group of population Whose members interbreed with one another successfully They are able to produce viable fertile offspring But cannot interbreed with member of other species Capable of genetic exchange share a common gene pool Limitations: 1. Not applicable to asexually reproducing species Examples: protists, fungi, plants (banana) bacteria 2. Cannot be applied to extinct organism represented only by fossils. 3. Difficult to apply the biological species to 2 populations that are geographically segregated. Example: Four phenotypically distinct populations of deer mouse are geographically isolated and are subspecies Presence of subspecies, or is this: Speciation (evolution) in action Will the species diverge or converge? Reproductive barriers: - Any factors that impede 2 species from producing fertile hybrids - contributes to reproductive isolation 2 major mechanism of reproductive isolation: 1. Prezygotic before the zygote is formed 2. Postzygotic after fertilization A. PREZYGOTIC 1. Habitat isolation - 2 species can live in the same area but may occupy different habitat Ex. Parasites living on different hosts will not have a chance to mate; one lives in water, the other on land. 2. Temporal Isolation

c) Directional : selective against

Speciation
2 patterns of evolutionary change: 1. Anagenesis change in one lineage 2. Cladogenesis Change in 2 or more lineage (branching) Anagenesis : transformation of an unbranched lineage To a state different enough form the ancestral population Justifies naming it a new species Cladogenesis: branching evolution The budding of one or more species from a parent species that continuous to exists More important than anagenesis in the history of life: a) More common b) Can promote biological diversity What is often observed is a lack of independent evolution

Six concepts of Species

1. 2. 3. 4. 5. 6. Biological species concept Morphological Recognition Cohesion Ecological Evolutionary

- 2 species breed at different times of the day, seasons or years, e.g. Brown trout (fall); rainbow trout (spring) 3. Behavioral isolation Difference in certain behavior -leads to failure in having a successful mating - refers to special signals and elaborate reproductive behavior unique to each species Ex. Mating calls, signals, songs, displays Pheromones 4. Mechanical Isolation - Anatomical incompatibility prevents sperm transfer of zygote formation Ex. a. Special appendages in insects b. Floral anatomy adapted to a specific pollinator 5. Gametic isolation - Different species lack gamete recognition -Due to specific molecules on the surface of the egg or sperm of different species.

Increase genetic variation occasional hybridization does not erase the boundary between corn and teosinte.

BIOGEOGRAPHY OF SPECIATION Modes of Speciation: 1. Allopatric 2. Parapatric 3. Sympatric A. Allopatric Speciation Due to a geographic barrier Species have separate ranges a. Become sympatric again and interbreed. Speciation did not occur. b. Become sympatric but did not interbreed B. Sympatric Speciation New species form within the range of the parent population Reproductive isolations occur without geographical isolation Mechanism: 1. Autopolyploidy results from a single species 2. Allopolyploidy hybrid results from 2 or more different species AUTOPOLYPLOIDY SPECIATION ALLOPOLYPLOIDY SPECIATION Triploid Crops: Bannana, apple, ginger, watermelon,

B. POSTZYGOTIC Prevent development of viable fertile hybrid 1. Hybrid inviablity - Development of the hybrid (embryo, fetus) is aborted at some stage 2. Hybrid Sterility - Offspring may be viable but sterile - Gene cannot flow from one gene pool to another - Major Cause: Chromosomes of the 2 parents differ in number and structure Ex. Mule: from donkey and horse 3. Hybrid Breakdown - First generation offsprings are viable and fertile But when hybrid mate with one another offspring are feeble or sterile C. INTROGRESSION The transplantation of alleles between species Ex. Corn (Zea mays) contains some alleles traceable to the closely related grass teosinte (Zea mexican) Occurs when 2 species hybridize and a small number of the hybrids manage to cross with corn plants.

citrus.

Tetraploid crops: Macaroni, wheat, maize, cotton, potato, cabbage, leek, tobacco, peanut Hexaploid Crops: Chrysanthemum, bread wheat, oat, kiwifruit Octaploid crops: strawberry, dahlia, sugar cane

Polyploidy in Vertebrates Parthenogenetic doubling of egg cell, no fertilization Peripatry varieties differ at the edge of the range becomes species similarly to allopatry. Parapatry variation evolves at the edge of the range become species similarly to sympathy. Allopatric divergence Sympatric polyploidy Parapatry difference in geographical ranges - Population has adjacent but not overlapping geographic ranges Population come in contact and meet over a hybrid zone Subspecies or geographic range (synonyms zoological taxonomy, plant taxonomy ecotypes) Recognizability distinct populations some can interbreed at hybrid zone Ex. Ratsnake (Elaphe obsoleta) has 5 species - Differ in a pattern and color - interbreed where the ranges meet