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What is a population? Review of Genetics Terminology A group of individuals (that can and do interbreed) living in a distinct geographic area Gene – sequence of DNA that codes Microevolution for a particular trait Evolution on a Allele – alternative versions of a gene small scale (within Genotype – combination of alleles a single species or present in the individual’s DNA population) Phenotype – the physical characteristics Change in gene resulting from the genotype frequency within a Gene pool – all the alleles for a What is a species? population gene that exist in a population group of individuals Mutation – random change in DNA o A that actually or sequence within an individual potentially interbreed in nature o The biggest gene pool possible under natural conditions Macroevolution Evolution on a grand scale (above the species level) Large-scale history of life Genes and evolution Only traits controlled by genes can be affected by natural selection. Genetic Variation • Necessary for evolution • Sources: • Mutations (random changes in genes) • Gene flow (movement of genes from one population to another of the same species) • Sexual Reproduction (creates new combinations of alleles within a population) Gene Frequency The number of times an allele occurs in a gene pool compared with the number of times other alleles occur EVOLUTION = CHANGE IN GENE FREQUENCIES! What causes a mutation? DNA fails to copy accurately during replication Damage to DNA from mutagen Types of gene mutations o Somatic mutation o Occurs in a non-reproductive cell o Not hereditary o Germ-line mutation o Occurs in a reproductive cell (gamete) o Heritable! o Drives evolution Potential effects of germ-line mutations No change in phenotype Small change in phenotype Major change in phenotype Mutation o Changes in DNA that affect phenotype o Example: Some beetles with “green genes” randomly mutate to “brown genes” Migration (Gene Flow) Movement of genes from one population to another population of the same species Genetic Drift Allele frequencies fluctuate unpredictably from one generation to the next Causes: First Generation Second Generation Random “shuffling” of alleles from one generation to the next Beetles example: 1st Generation: 75% Green 25% Brown 2nd Generation: 71% Green 29% Brown 75% 25% 71% 29% Genetic Drift Allele frequencies fluctuate unpredictably from one generation to the next Causes: Random “shuffling” of alleles from one generation to the next Bottleneck Effect: Sudden change in environment 1. Initial population has equal drastically reduces population frequencies of red and yellow “alleles” and alters allele frequencies 2. Chance event greatly reduces the size of the population 3. Surviving individuals have “allele” frequencies different from those of the original population 4. Resulting population has far more red “alleles” than yellow 1 2 3 4 Genetic Drift Allele frequencies fluctuate unpredictably from one generation to the next Causes: Random “shuffling” of alleles from one generation to the next Bottleneck Effect: Sudden change in environment drastically reduces population and alters allele frequencies Cheetahs: so close that European bison: entire living population today descends from only 12 individuals. skin grafts do not trigger an immune response Elephant seals: only 30 in the 1890s; 10’s of thousands today Genetic Drift Allele frequencies fluctuate unpredictably from one generation to the next Causes: Random “shuffling” of alleles from one generation to the next Bottleneck Effect Founder Effect: A few individuals from a population start a new population with a different allele frequency than the original population 1. Several individuals become isolated from a larger population and… 2. …establish a new population. 3. The new population’s gene pool is not reflective of the source population. 1 2 3 Genetic Drift Allele frequencies fluctuate unpredictably from one generation to the next Causes: Random “shuffling” of alleles from one generation to the next Bottleneck Effect Founder Effect: A few individuals from a population start a new population with a different allele frequency than the original population Sample of Original Population Founding Population A Descendants of A Founding Population B Descendants of B 1. Several individuals become isolated from a larger population and… 2. …establish a new population. 3. The new population’s gene pool is not reflective of the source population. Natural Selection • Changes in the inherited characteristics of a population, which increase the species’ fitness in its environment • Example: Predators eat more of the green ones, so brownof is beetles more Original population likely togreen survive is predominantly and reproduce Remaining population Green beetles are more is predominantly brown visible against tree bark than brown ones. Fitness Ability of an individual to survive and reproduce in its specific environment Natural Selection • Changes in the inherited characteristics of a population, which increase the species’ fitness in its environment • Example: Natural Selection • Changes in the inherited characteristics of a population, which increase the species’ fitness in its environment • Example: Natural Selection • Changes in the inherited characteristics of a population, which increase the species’ fitness in its environment • Example: Natural Selection • Changes in the inherited characteristics of a population, which increase the species’ fitness in its environment • Example: Natural Selection • Changes in the inherited characteristics of a population, which increase the species’ fitness in its environment • Example: 1850 1900 Natural Selection • Changes in the inherited characteristics of a population, which increase the species’ fitness in its environment Sexual Selection: • Non-random mating within a population Sexual Dimorphism Males and females of a species differ significantly in appearance Natural Selection • Changes in the inherited characteristics of a population, which increase the species’ fitness in its environment Sexual Selection: Sexual Dimorphism Males and females of a species differ significantly in appearance • Non-random mating within a population • Types of sexual selection: • Intrasexual selection • Competition within the same sex for mates of the opposite sex • Favors traits that deal with conflict between individuals of the same sex Natural Selection • Changes in the inherited characteristics of a population, which increase the species’ fitness in its environment Sexual Selection: Sexual Dimorphism Males and females of a species differ significantly in appearance • Non-random mating within a population • Types of sexual selection: • Intrasexual selection • Competition within the same sex for mates of the opposite sex • Favors traits that deal with conflict between individuals of the same sex Natural Selection • Changes in the inherited characteristics of a population, which increase the species’ fitness in its environment Sexual Selection: Sexual Dimorphism Males and females of a species differ significantly in appearance • Non-random mating within a population • Types of sexual selection: • Intrasexual selection • Competition within the same sex for mates of the opposite sex • Favors traits that deal with conflict between individuals of the same sex • Intersexual selection • Individuals of one sex (usually female) are selective when choosing a mate • Favors traits that enhance attractiveness to the opposite sex, such as plumes, wattles, bright color, songs, symmetry, etc. Natural Selection • Changes in the inherited characteristics of a population, which increase the species’ fitness in its environment Sexual Dimorphism Males and females of a species differ significantly in appearance Sexual Selection: Cauliflower • Non-random mating within a population • Types of sexual selection: • Intrasexual selection • Competition within the same sex for mates of the opposite sex • Favors traits that deal with conflict between individuals of the same sex • Intersexual selection Wild • Individuals of one sex (usually female) Mustard are selective when choosing a mate • Favors traits that enhance attractiveness to the opposite sex, such as plumes, wattles, bright color, songs, symmetry, etc. Brussels Sprouts Flower clusters Lateral buds Cabbage Terminal buds Kale Broccoli Artificial Selection: • Selective breeding • Natural variations exist within a species, but humans select which organisms get to reproduce Stems Kohlrabi Leaves Stems & flowers Stabilizing Selection Trend toward reducing phenotypic variation and maintenance of the status quo Individuals with the average phenotype (center) are selected Examples: Tuatara reptiles – very little change over long periods of time Birth weight of humans – too small or too large tend to have lower survival rates Selection against both extremes Very low birth weight babies are less likely to survive Extremely large babies may not survive birth (may also cause death of mother during delivery) Original population ~7.5 lbs Average weight Population after selection Directional Selection Trend toward one extreme and away from the other extreme, such that the population as a whole tends to change in a similar fashion Individuals with one phenotype (at one end) are selected Examples: Peppered moths, insecticide resistance, antibiotic resistance Selection against one extreme Original population Pre-industrial moths were light-colored Population after selection Post-industrial moths were dark-colored Disruptive Selection Two or more phenotypes survive and reproduce more successfully than intermediate forms Individuals with one of two phenotypes (both ends) are selected Example: African Swallowtail Butterfly Selection against the average Female mimics an unpalatable species Original population Population after selection Male or female is palatable and does not mimic any unpalatable species Female mimics a different unpalatable species Stabilizing selection Individuals with the average phenotype (center) are selected Original population of brown mice Directional selection Individuals with one phenotype (one end) are selected Disruptive selection Individuals with one of two phenotypes (both ends) selected Original population Phenotypes (fur color) Population after selection Wide variation in fur color Medium colors selected Medium color selected Stabilizing Selection Darker colors selected Directional Selection against Disruptive Selection Macroevolution Evolution above the species level Overall trends and transformations in evolution Speciation • Development of new species. • Over long periods of time, natural selection leads to the accumulation of changes that differentiate groups from one another. Patterns in macroevolution 1. Stasis Lineages that change very little over long periods of time (ex: coelocanths) 2. Character change Characteristics of lineages can change quickly or slowly over time Characteristics may change in a consistent manner or may progress and regress over time 3. Lineage-splitting (speciation) Speciation within a lineage can happen often or infrequently 4. Extinction May occur frequently or rarely in a lineage Simultaneous extinctions across many lineages are known as “mass extinctions” What distinguishes one species from another? Morphological Species Concept: Characterizes a species by its body shape, size, and other structural features Applies to asexual and sexual organisms Useful without information on gene flow Often used in paleontology, since little information may be known about mating abilities of fossilized extinct organisms. Phylogenetic Species Concept: Set of organisms with a unique genetic history Relatedness determined by: Comparison of physical characteristics Molecular similarities (DNA) Ecological Species Concept: Focuses on differences in ecological niche rather than appearance (i.e. food source) Biological Species Concept: Groups of actually or potentially interbreeding natural populations Produce fertile offspring Reproductively isolated from other groups Commonly accepted as a basic working definition of species (we’ll use this one!) Therefore… …two populations are separate species if they cannot (or do not) successfully interbreed in nature. What factors may prevent successful interbreeding between populations (aka reproductive isolation)? Pre-zygotic Barriers: Factors that may inhibit: Mating between individuals Fertilization of gametes Post-zygotic Barriers: Factors that may inhibit: Survival of embryo or offspring Fertility of surviving offspring Pre-zygotic Barriers 1. Geographic isolation – populations are separated by a physical boundary (i.e. river) What factors may prevent successful interbreeding between populations (aka reproductive isolation)? Geographic Isolation What factors may prevent successful interbreeding between populations (aka reproductive isolation)? Pre-zygotic Barriers: Factors that may inhibit: Mating between individuals Fertilization of gametes Post-zygotic Barriers: Factors that may inhibit: Survival of embryo or offspring Fertility of surviving offspring Pre-zygotic Barriers 1. Geographic isolation – populations are separated by a physical boundary (i.e. river) 2. Ecological isolation – populations live in the same area, but do not meet due to differences in ecology (i.e. different pollinators or food sources) What factors may prevent successful interbreeding between populations (aka reproductive isolation)? Ecological Isolation (different pollinators) What factors may prevent successful interbreeding between populations (aka reproductive isolation)? Different habitats: •Open woods •Beech trees •Alders •Conifer woods •Willowy thickets Ecological Isolation What factors may prevent successful interbreeding between populations (aka reproductive isolation)? Pre-zygotic Barriers: Factors that may inhibit: Mating between individuals Fertilization of gametes Post-zygotic Barriers: Factors that may inhibit: Survival of embryo or offspring Fertility of surviving offspring Pre-zygotic Barriers 1. Geographic isolation – populations are separated by a physical boundary (i.e. river) 2. Ecological isolation – populations live in the same area, but do not meet due to differences in ecology (i.e. different pollinators or food sources) 3. Temporal isolation – mating or flowering occurs at different seasons or time of day What factors may prevent successful interbreeding between populations (aka reproductive isolation)? Temporal Isolation What factors may prevent successful interbreeding between populations (aka reproductive isolation)? (breeds in September/October) (breeds in March/April) Temporal Isolation What factors may prevent successful interbreeding between populations (aka reproductive isolation)? Pre-zygotic Barriers: Factors that may inhibit: Mating between individuals Fertilization of gametes Post-zygotic Barriers: Factors that may inhibit: Survival of embryo or offspring Fertility of surviving offspring Pre-zygotic Barriers 1. Geographic isolation – populations are separated by a physical boundary (i.e. river) 2. Ecological isolation – populations live in the same area, but do not meet due to differences in ecology (i.e. different pollinators or food sources) 3. Temporal isolation – mating or flowering occurs at different seasons or time of day 4. Behavioral isolation – little or no sexual attraction between males and females What factors may prevent successful interbreeding between populations (aka reproductive isolation)? Behavioral Isolation (different mating songs) What factors may prevent successful interbreeding between populations (aka reproductive isolation)? Pre-zygotic Barriers: Factors that may inhibit: Mating between individuals Fertilization of gametes Post-zygotic Barriers: Factors that may inhibit: Survival of embryo or offspring Fertility of surviving offspring Pre-zygotic Barriers 1. Geographic isolation – populations are separated by a physical boundary (i.e. river) 2. Ecological isolation – populations live in the same area, but do not meet due to differences in ecology (i.e. different pollinators or food sources) 3. Temporal isolation – mating or flowering occurs at different seasons or time of day 4. Behavioral isolation – little or no sexual attraction between males and females 5. Mechanical isolation – anatomical differences prevent exchange of gametes What factors may prevent successful interbreeding between populations (aka reproductive isolation)? Mechanical Isolation What factors may prevent successful interbreeding between populations (aka reproductive isolation)? Pre-zygotic Barriers: Factors that may inhibit: Mating between individuals Fertilization of gametes Post-zygotic Barriers: Factors that may inhibit: Survival of embryo or offspring Fertility of surviving offspring Pre-zygotic Barriers 1. Geographic isolation – populations are separated by a physical boundary (i.e. river) 2. Ecological isolation – populations live in the same area, but do not meet due to differences in ecology (i.e. different pollinators or food sources) 3. Temporal isolation – mating or flowering occurs at different seasons or time of day 4. Behavioral isolation – little or no sexual attraction between males and females 5. Mechanical isolation – anatomical differences prevent exchange of gametes 6. Gametic isolation – female and male gametes fail to unite in fertilization What factors may prevent successful interbreeding between populations (aka reproductive isolation)? Gametic Isolation What factors may prevent successful interbreeding between populations (aka reproductive isolation)? Pre-zygotic Barriers: Factors that may inhibit: Mating between individuals Fertilization of gametes Post-zygotic Barriers: Factors that may inhibit: Survival of embryo or offspring Fertility of surviving offspring Pre-zygotic Barriers 1. Geographic isolation – populations are separated by a physical boundary (i.e. river) 2. Ecological isolation – populations live in the same area, but do not meet due to differences in ecology (i.e. different pollinators or food sources) 3. Temporal isolation – mating or flowering occurs at different seasons or time of day 4. Behavioral isolation – little or no sexual attraction between males and females 5. Mechanical isolation – anatomical differences prevent exchange of gametes 6. Gametic isolation – female and male gametes fail to unite in fertilization Post-zygotic Barriers 1. Hybrid inviability – hybrid zygotes fail to develop or fail to reach sexual maturity What factors may prevent successful interbreeding between populations (aka reproductive isolation)? Hybrid Inviability What factors may prevent successful interbreeding between populations (aka reproductive isolation)? Pre-zygotic Barriers: Factors that may inhibit: Mating between individuals Fertilization of gametes Post-zygotic Barriers: Factors that may inhibit: Survival of embryo or offspring Fertility of surviving offspring Pre-zygotic Barriers 1. Geographic isolation – populations are separated by a physical boundary (i.e. river) 2. Ecological isolation – populations live in the same area, but do not meet due to differences in ecology (i.e. different pollinators or food sources) 3. Temporal isolation – mating or flowering occurs at different seasons or time of day 4. Behavioral isolation – little or no sexual attraction between males and females 5. Mechanical isolation – anatomical differences prevent exchange of gametes 6. Gametic isolation – female and male gametes fail to unite in fertilization Post-zygotic Barriers 1. Hybrid inviability – hybrid zygotes fail to develop or fail to reach sexual maturity 2. Hybrid infertility – hybrids fail to produce functional gametes What factors may prevent successful interbreeding between populations (aka reproductive isolation)? Hybrid Infertility What factors may prevent successful interbreeding between populations (aka reproductive isolation)? Pre-zygotic Barriers: Factors that may inhibit: Mating between individuals Fertilization of gametes Post-zygotic Barriers: Factors that may inhibit: Survival of embryo or offspring Fertility of surviving offspring Pre-zygotic Barriers 1. Geographic isolation – populations are separated by a physical boundary (i.e. river) 2. Ecological isolation – populations live in the same area, but do not meet due to differences in ecology (i.e. different pollinators or food sources) 3. Temporal isolation – mating or flowering occurs at different seasons or time of day 4. Behavioral isolation – little or no sexual attraction between males and females 5. Mechanical isolation – anatomical differences prevent exchange of gametes 6. Gametic isolation – female and male gametes fail to unite in fertilization Post-zygotic Barriers 1. Hybrid inviability – hybrid zygotes fail to develop or fail to reach sexual maturity 2. Hybrid infertility – hybrids fail to produce functional gametes 3. Hybrid breakdown – 1st generation hybrids can reproduce, but 2nd cannot What factors may prevent successful interbreeding between populations (aka reproductive isolation)? F1 hybrids are viable F2 fails to germinate Hybrid Breakdown The hybrid produced by a lion and a tigress (known as a "liger") is usually larger and stronger than either of its parents, whereas the reciprocal cross produces a hybrid (the "tigon") that tends to be smaller than either of its parents (Gray 1972, McCarthy, in prep.). The former cross, then, is an example of positive heterosis, in which the hybrid exceeds the range of variation exhibited by its parents, while the latter is an example of negative heterosis, in which the hybrid falls below the range of parental variation with respect to a given trait. Phyletic Gradualism Speciation caused by small adaptations accumulating throughout the history of a species in response to new environmental and biological selective pressures Changes are smooth, steady, and incremental rate (on a geological timescale) Changes are difficult to notice over short periods of time No clear line of demarcation between an ancestral species and a descendant species unless a splitting event occurs or the graduallychanging lineage is divided arbitrarily. Problem: transitional fossils? Punctuated Equilibrium Changes come in spurts Long periods of stasis followed by very brief periods of great change that result in abrupt lineage-splitting (speciation) Changes may be due to random mutations or sudden changes in the environment Explains lack of transitional fossils Punctuated Gradualism Long periods of gradual change may be interrupted by brief periods of rapid change without leading to lineage-branching Punctuated Equilibrium Phyletic Gradualism Specialists May thrive only in a narrow range of environmental conditions May have a limited diet Examples: Panda: feed primarily on bamboo Koala: diet consists almost exclusively of eucalyptus leaves Advantages of specializing: More effective at competing in its specific niche Tend to thrive in stable environments Disadvantages of specializing: Populations may be limited due to limited suitable environments More likely to go extinct when conditions change Generalists Often able to adapt to a wide range of environments Diet may be varied (omnivores are often generalists) Frequently adapt well to urban environments Examples: Opossums Raccoons Advantages of being a generalist: May occupy much wider ranges, allowing much larger populations More likely to adapt well to changes in environment Disadvantage of being a generalist: May be less able to compete with specialists during periods of stable environment