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Natural selection & adaptation

Habitats

Fisheries Management

Changing World

Ecology & Evolution

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Darwin’s dangerous idea

Identified a process that could explain the patterns of diversity on Earth and (even though he never spoke of speciation per se) could provide the ‘origin of species’ (the mystery of mysteries as he called it).

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Darwin’s dangerous idea

Identified a process that could explain the patterns of diversity on Earth and (even though he never spoke of speciation per se) could provide the ‘origin of species’ (the mystery of mysteries as he called it).

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Contentious then (and now) because the other proposed mechanism involves supernatural processes

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Low rate of acceptance of evolution in the US

Response to the following true or false question: “human beings, as we know them, developed from earlier species of animals”

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Researchers working in this area have identified lack of understanding of natural selection as a process as a source of ‘not sure’ and (at least some) of the ‘false’ responses.

Miller et al. 2006

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Signatures of evolution

# 1: Major changes in traits come through gradual modification of existing forms

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#2: Major change (e.g. complete speciation, reproductive isolation) takes thousands of generations, though important changes in trait size, shape, etc… (microevolution) can occur in only a few generations

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The process; evolution by natural selection

  1. The advantage of individuals of a given species over other members with specific regards to survival and reproduction (ecologist’s definition)
  2. The accumulation of beneficial alleles (i.e. gene variants. Recall from intro biology A and a; geneticist’s definition)
  3. The non-random sorting of random variation (a third reasonable definition)

Some complementary definitions of natural selection

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Fitness!

“Something that everyone knows but can’t define precisely”- Steve Stearns, famous evolutionary biologist

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Fitness!

“Something that everyone knows but can’t define precisely”- Steve Stearns, famous evolutionary biologist

“…if the variation were of a beneficial nature, the original form would soon be supplanted by the modified form, through the survival of the fittest.” Darwin in the Origin

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Fitness!

“Something that everyone knows but can’t define precisely”- Steve Stearns, famous evolutionary biologist

“…if the variation were of a beneficial nature, the original form would soon be supplanted by the modified form, through the survival of the fittest.” Darwin in the Origin

(Darwinian ) fitness is thus a term that describes the ability of an individual to survive and reproduce

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Population fitness is a measure of population growth potential (or growth rate, λ). Populations with high fitness are comprised by individuals with high Darwinian fitness

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The logic behind natural selection

Premise 1: populations can produce many more individuals than the environment can support, therefore there is competition for limited resources, “struggle for existence”

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The logic behind natural selection

Premise 1: populations can produce many more individuals than the environment can support, therefore there is competition for limited resources, “struggle for existence”

Premise 2: some individuals have traits that convey advantage in the struggle for resources and those individuals are more likely to survive and reproduce and produce more children than others, “survival of the fittest”

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The logic behind natural selection

Premise 1: populations can produce many more individuals than the environment can support, therefore there is competition for limited resources, “struggle for existence”

Premise 2: some individuals have traits that convey advantage in the struggle for resources and those individuals are more likely to survive and reproduce and produce more children than others, “survival of the fittest”

Premise 3: if traits that convey advantage have a heritable (i.e. genetic) component they will be propagated in future generations and individuals will become adapted to their local environments, “adaptive evolution”.

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The logic behind natural selection

Premise 1: populations can produce many more individuals than the environment can support, therefore there is competition for limited resources, “struggle for existence”

Premise 2: some individuals have traits that convey advantage in the struggle for resources and those individuals are more likely to survive and reproduce and produce more children than others, “survival of the fittest”

Premise 3: if traits that convey advantage have a heritable (i.e. genetic) component they will be propagated in future generations and individuals will become adapted to their local environments, “adaptive evolution”.

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R = h2S (breeders equation)

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In other words…

Premise 1: Competition

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Premise 2: Trait variations allow some individuals to compete better than others (and thus have higher prob. of survival and reproduction

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Premise 3: if random variation is genetic, then adaptive evolution occurs

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A first example…brewer’s yeast

Fermentation occurs when yeast converts the glucose (sugar) in the grain mixture (wort) into ethyl alcohol and CO2

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Fermentation ceases due to depleted sugar and toxicity of alcohol to yeast (the booze in your beer is a deadly by- product to the yeast)

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Let’s provide an explanation for increased alcohol tolerance in yeast

Inspiring brew master PJ Clark is working on mastering his IPA recipe. With the goal of saving money he harvests the surviving yeast from his beer at the end of the fermentation step and uses those yeast to seed his next batch of brew. He measures the alcohol content (inferred through specific gravity) of each batch. The results are tabulated below:

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Batch

Final* Specific gravity (P)

Alcohol % by volume

1

22

4.89

2

20

6.06

3

18

7.22

4

15

8.93

5

14

9.50

6

12

10.6

Original SG = 30

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Let’s take a few minutes in groups

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Let’s provide an explanation for increased alcohol tolerance in yeast

Batch

Final* Specific gravity (P)

Alcohol % by volume

1

22

4.89

2

20

6.06

3

18

7.22

4

15

8.93

5

14

9.50

6

12

10.6

Thoughts?

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Group 1: Yeast genetically changes – outliving other yeast that aren’t as viable

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Group 2: Yeast is evolving from batch to batch – getting stronger. Using more of the sugars to release more alcohol

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Group 3: same yeast is producing enzymes better over time

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Group 4: Stronger alcohol tolerance – leftover yeast

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Group 5: Genetically advantageous to become more resistant.

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A potential response

The population of yeast used by PJ Clark had existing genetic variation in the resistance of yeast cells to different levels of alcohol. Cells that were more tolerant to alcohol survived whereas more susceptible cells died. The resistance to alcohol has a genetic basis and therefore resistance increases in subsequent generations of yeast.

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Note: Increasing alcohol represents a change in the environment for yeast, making some individual maladapted to those conditions. Individuals with traits that allow them to survive in the new environment are favored.

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The process of change is adaptive evolution by natural selection

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The mechanism of selection was susceptibility of certain enzymes in the yeast to alcohol.

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Process vs. mechanism

Natural selection is the process of change.

The mechanism favoring some individuals is ecological, physiological, behavioral, etc…

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Predation by piscivorous fish eating the most conspicuously colored prey is an ecological mechanism of selection

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What was the mechanism of selection in the yeast example?

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Provide an explanation…

the short fin mako can burst speed to over 50 km/h, but its closest common ancestor can burst speed to only 30 km/h

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Sources of variation

Two major sources of variation that is important for evolution of populations (one is contentious!)

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  • Random mutations. Selection can favor mutations that exist in the population (standing variation) or from de novo mutations (variation that arises anew). This is the default standard explanation for variation in populations.

For example: ATTCGCCCGGAATTAAGAGTGAGG

is changed to: ATTCGCCCGAAATTAAGAGTGAGG

This code gives the “blue print” for the protein dystophin

‘normal’ protein shape

‘normal’ protein shape

  • Genetic Drift. variation in the relative frequency of different genotypes in a small population, owing to the chance disappearance of particular genes as individuals die or do not reproduce.

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Hopeful mutants

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Other types of variation

  • Phenotypic plasticity. Traits that are determined by environmental factors (i.e controlled more by the environment than genetics). The role of plasticity in the evolution of populations remains hotly debated! This is a reminder that selection can be very strong on traits that have little or no genetic basis.

Former arm-wrestling world champion!

Raised without predator

Raised with predator

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Protected bay

Exposed bay

Proc. R. Soc. B (2008) 275, 1081–1087

doi:10.1098/rspb.2007.1760

Published online 5 February 2008

Adaptive evolution or plasticity?

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Let’s take a few minutes in groups

Provide a hypothesis for this observed pattern, and a potential way to test to see if you were right

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This is what Chris Neufeld did!

Interpret the results

Protected

Exposed

Protected

Exposed

Growth Environment

Growth Environment

Came from: Exposed Protected

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FTT Natural or artificial (unnatural)

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Natural or artificial (unnatural)

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Natural or artificial (unnatural)

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Natural or artificial (unnatural)

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Natural or artificial (unnatural)

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Natural or artificial (unnatural)

Natural or artificial (unnatural)

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Natural or artificial (unnatural)

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Natural or artificial (unnatural)

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Natural or artificial (unnatural)

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Natural or artificial (unnatural)

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Natural or artificial (unnatural)

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Evolution of blindness in cave fishes

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Evolution of blindness in cave fishes

Some facts

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Cave form: Blind (no eyes), no pigment, heightened brain activity in region that controls olfaction (sense of smell) and loss of brain region controlling vision. Very efficient at building fat reserves. Non-schooling behavior.

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Surface form: Acute vision, heavily pigmented, relatively large brain area for vision and relatively little for olfaction. Less efficient at storing fat reserves. Occur in large schools.

Cave form

Surface form

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Provide an explanation

For the loss of eyes and pigmentation in cave-dwelling tetras, given that cave populations were colonized by surface dwelling individuals that had eyes and were pigmented.

Cave form

Surface form

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Let’s take a few minutes in groups (3-4 people…you decide)

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Thoughts?

For the loss of eyes and pigmentation in cave-dwelling tetras given that cave populations were colonized by surface dwelling individuals that had eyes and were pigmented.

Cave form

Surface form

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Propose a way to determine

Whether the lack of eyes and pigmentation in cave-dwellers is primarily the result of the environment or genetics?

Cave form

Surface form

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Let’s take a few minutes in groups (3-4 people…you decide)

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Ideas on how to disentangle genes from the environment?

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Common garden experiments

Logic: take groups of individuals that differ in traits of interest (in this case eyes and pigmentation) and let them grow and develop in the same environment. If surface forms maintain eyes (and pigment) and blind-forms remain blind (and lack pigment) then this is strong evidence for genetic control of these traits.

Genes or environment???

Same conditions

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Modified common garden experiments

Logic: Rear surface-dwellers and cave-dwellers in conditions that are similar to their own and each other’s environment and measure ‘performance’ (traits linked to ‘fitness’: growth, survival, reproductive output, etc…) in each environment.

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Hypothesis and predictions assuming adaptation: ???

Cave-like conditions

Surface-like conditions

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Reciprocal transplants

Move surface-dwellers to cave environments and cave-dwellers to surface environments and quantify survival, growth, or other fitness-linked traits.

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A much more realistic and powerful approach.

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And much more difficult to do logistically!

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Evolution of coloration in male guppies

Photos: A. Hendry

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Started with an observation in nature

Photos by A. Hendry & P. Bentzen

Below waterfalls

Above waterfalls

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Hypotheses?

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Streams in Trinidad below waterfalls are dangerous places (for guppies)

Rivulus hartii, killifish

Occurs with guppies above waterfalls. Eats guppies but only very small ones (gape-limited predator)

Occurs with guppies below waterfalls. Loves to eat adult guppies. Any guesses what sense this predator uses most for hunting?

Crenicichla alta, pike cichild

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Are predators driving the color patterns in guppies?

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How might you design an experiment to test for effect of predation on guppy color?��Two minutes alone…

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6

7

8

9

10

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Ideas??

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One (good) approach

John Endler

  1. Create artificial but realistic streams and stock them with a population of guppies with similar starting coloration patterns
  2. Introduce a strong predator to some of the artificial streams (high-predation treatment), a gape-limited predator (low-predation treatment) and leave the others predator free (control)
  3. Record the coloration patterns in the different streams through time
  4. Also…take guppies from high-predation habitats and transplant them above waterfalls into low-predation habitats

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Results from artificial streams

Endler 1980

Founding generation from Trinidad

Predators introduced

High predation habitats

No predation habitats

Low predation

~ 10 generations

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Predation leads to less colorful guppies

But why do guppies get more colorful in absence of predation??

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Sexual selection

The advantage of individuals over others of the same species with specific regards to reproduction

“Special means of defence may be given through means of sexual selection, as the mane to the lion, the shoulder-pad to the boar, and the hooked jaw to the male salmon…”

Darwin, the Origin

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Summary

  • Evolution = mutation, selection, genetic variation (heritability)
  • Fitness = the “currency of selection”
    • Survival, reproduction, growth
  • Ecology and evolution influence each other
  • Common garden + reciprocal transplant experiments can help differentiate ecologically-driven variation (plasticity) from genetically-driven variation (evolution)

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