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Mechanisms of Evolution
- Allelic frequency - Hardy-Weinberg - Polymorphism - Patterns of natural selection - Mechanisms of isolation - Rates of evolution Chapter 11 in text
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Individuals do not evolve –
Populations do. Individuals may be selected, and contribute differentially to the future gene pool. Gene mutations provide the genetic variability. Natural selection acts on the collective genes in the gene pool of a population. An individual has the genes he was born with (nothing will change that), but the frequency of a given allele in a population may change over time. This is evolution.
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Allelic frequency
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NB: We are seeing the frequency of these phenotypes
in the entire population… 3:1 assumes p = 0.5, q = 0.5
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- Hardy-Weinberg For a gene with two alleles, Hardy-Weinberg equation: p2 + 2pq + q2 = 1 The sums of the genotype frequencies equals 1... p + q = 1 The sum of the allelic frequencies = 1, too An example… 1 of every 375 African-Americans has sickle cell anemia. What proportion of this population has sickle cell trait? ss homozygous = 1/375, or = q2 ______ q = √ = 0.052, sooo p = 1 – = 0.948 Ss is 2pq, = 2(0.948)(0.052) = 0.099 or 1- (p2 + q2) = 1- ( ), = 1- ( ) = 0.099, or so, I rounded along the way.
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to study something, it is often helpful to know what it isn’t…
Hardy-Weinberg equilibrium: The Hardy-Weinberg equation will describe the allelic frequency in a population if no evolution is happening. Assumptions of such an equilibrium: 1. Extremely large population 2. No gene flow 3. No mutations 4. Random mating 5. No natural selection We will look at each of these in turn.
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1. Extremely large population
If a population is small, allelic frequency may change significantly through random chance or sampling error. Picture 6 birds of one species arriving on an island, If they have the same color distribution as the parent population, but the two lightest happen to have fewer offspring, the subsequent population will be darker. This is called genetic drift.
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1. Extremely large population (cont.)
Let’s say these first six birds are all equally successful in breeding, but they happened to be a darker group to begin with, compared to the parent population. This is called the founder effect. faculty.clintoncc.suny.edu/.../evolutio.htm
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1. Extremely large population (cont.)
Let’s say the original, diverse morphs populated the island, but a hurricane took out all but a few, which happened to be lighter ones… This is called the bottleneck effect. These three are interconnected (small founder group is more subject to drift…)
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2. No gene flow This means no significant immigration or emigration… No gene flow into or out of the population. Naturally, adjacent populations may experience integration occasionally→ This could change the allelic frequencies.
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3. No mutations We have discussed gene and chromosome mutations: - These tend to occur at a predictable frequency. - They may be advantageous (adaptive), disabling (maladaptive), or inconsequential (silent or neutral variation). Without the underlying variety that mutations enable, no evolution would be possible.
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Darwin was one of the first to propose sexual selection,
4. Random mating Darwin was one of the first to propose sexual selection, because the displays otherwise seem maladaptive - (wastes of energy and unnecessary exposure). Intrasexual selection: competition between members of the same gender, like elk rutting displays. Intersexual selection: choosing from one of the opposite gender, like peahens and peacocks (sexual dimorphism). - The displays may reflect overall health and vigor, apart from simply the apparent foppery. - Note: doesn’t apply to asexually reproducing organisms…
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5. No natural selection Natural selection (for an adaptive trait) is the poster child of evolution. The key is that those traits that best allow individuals to survive to adulthood and successfully reproduce, will be passed along to the next generation at the highest frequency. positive selection Less favorable traits will become less frequent. negative selection
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5. No natural selection (cont.)
Sometimes an advantage seems obvious (disease resistance), sometimes less so (altruism).
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So, does Hardy-Weinberg equilibrium ever apply?
- Some organisms are preserved relatively intact for huge amounts of time, as evidenced in the fossil record. - Probably many species have come and gone too quickly to be preserved. If the Hardy-Weinberg equation is found to not describe a population, it is time to look at what mechanisms of evolution are at play.
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caused by different alleles for different genes.
- Polymorphism Within a population there is a range of forms (morphs) for a variety of traits, caused by different alleles for different genes. Some traits are polygenic, and exhibit a continuum of morphology. Why isn’t it better to have one trait, or the other? Two (or more) alleles cooperating is sometimes better than either one alone. e.g. HsHs → sickle-cell anemia HHs → OK (heterozygous advantage) HH → more susceptible to malaria
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Transient polymorphism:
One version is favored sometimes, the other at other times, e.g. industrial melanism and the peppered moth. Note: this is microevolution, not speciation, but provides a testable example of natural selection.
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- Patterns of natural selection
stabilizing selection: The central-most morph is most successful, and extreme forms are reduced. Results in fine-tuned, but potentially fragile species. (Why?) disruptive selection: The central form is less adaptive, and the population splits into two. Due to competition, loss of original resource... Easy step to speciation. directional selection: One extreme is favored over the other. Generally leads to speciation only if two populations are already separated.
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divergent evolution: The result of divergent selection, in which one species faces selective pressure for two different niches: Recognized by presence of homologous structures. e.g. Australian apple maggot fly: originated from the Australian hawthorn maggot fly, but when apples were brought in, the new food source was optimized... Now two distinct species exist.
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adaptive radiation is a specific case of divergent evolution, in which many niches are available for exploitation e.g. the post-Mesozoic explosion of mammals after mass extinction of dinosaurs
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Adaptive radiation also happens when organisms colonize a new island grouping… This resembles the process of the finches in the Galapagos islands.
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convergent evolution:
A specific type of directional selection, in which two species are long separated, but selected towards a common adaptation: Often recognized by the presence of analogous structures.
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- Mechanisms of isolation
In order for actual speciation to occur, there has to be some sort of boundary preventing the mixing of genes… Reproductive isolation… self explanatory. - prevention of successful interbreeding, ….somehow. Two main groupings: 1. Geographic (“allopatric”) – Organisms are physically separated. This could be across an ocean, or a stream, or a spit of land, or adjacent puddles. 2. Sympatric – (“same country”) Various mechanisms that isolate neighbors…
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1. Geographic isolation Your text has an example of squirrels and the Galapagos finches.
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2. Sympatric isolation habitat isolation: different use of an ecosystem. e.g. water vs. land garter snakes behavioral isolation: courtship, communication e.g. blue-footed boobies, Pacific killer whales temporal isolation: time of day, year, cycles… e.g. E spotted skunk (winter) vs. W (summer) mechanical isolation: structural incompatibility… 2 monkey flowers attract different pollinators gametic isolation: repro. tract or gamete surface sea urchins
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- Rates of evolution Microevolution by selection or genetic drift can happen in the course of a few generations. Speciation by polyploidy happens in a single generation. More commonly, the accumulation of adaptive mutations can be very slow (thousands of years? tens of thousands?), and if an organism is well adapted, there may be little change in allelic frequency over millions of years. HOWEVER, after a disruption, there is generally astonishingly rapid evolution as newly freed or created niches are filled, depending on existent mutations. - Punctuated equilibrium
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If a given dominant trait is expressed
in 450 of every 1000 individuals in a population, what is the expected frequency of heterozygotes? Distinguish among bottleneck effect, founder effect, and genetic drift. Propose illustrations for the three types of selection - stabilizing, directional, and disruptive. What is Hardy-Weinberg equilibrium? Describe the evolutionary factors that contributed to the speciation of “Darwin’s” finches. What is evolution?
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Benchmark SC.912.L.15.13 Describe the conditions required for natural selection, including overproduction of offspring, inherited variation, and the struggle to survive, which results in differential reproductive success.
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Life on Earth is very diverse, and even within species
Enduring Understandings Life on Earth is very diverse, and even within species there is great diversity. Essential Questions What mechanisms have allowed for diversity in organisms?
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gene pool sexual dimorphism evolution natural selection Hardy-Weinberg equation positive selection p2 + 2pq + q2 = 1 negative selection p + q = 1 polymorphism Hardy-Weinberg equilibrium heterozygous advantage 1. Extremely large population stabilizing selection genetic drift disruptive selection founder effect directional selection bottleneck effect divergent evolution 2. No gene flow adaptive radiation 3. No mutations convergent evolution 4. Random mating reproductive isolation 5. No natural selection geographic isolation sexual selection sympatric isolation intrasexual selection behavioral isolation intersexual selection temporal isolation speciation punctuated equilibrium mutation genetic recombination diversity extinction allele gene pool genetic drift gene migration/gene flow Hardy-Weinberg principle
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