Testing the Hypothesis of

Natural Selection continued

First, a look at various adaptations
discussed by Coyne (Chapter 5)
 Another Drought example: testing
hypotheses of natural selection
Do bird beak-sizes respond to selection by
changing morphology?
Another Example: Beak shape in the
Galapagos Finches
All species are close relatives
Differ with respect to beak size &
shape
 Different beak shapes, different diets
Large Ground Finch: seeds Sharp-beaked Ground Finches
 Different beak shapes, different diets
Vegetarian finch: leaves and fruits Warbler finches: arthropods
Peter and Rosemary Grant: Studying
Darwin’s finches since 1973
• Geospiza fortis (medium ground finch) on
Daphne major
 Grants’ studies
• Daphne major: Warm, wet – Jan – May.
Cool, dry – June – Dec.
• ALL medium ground finches have been
banded since 1980.
• Primarily seed eaters.
• Interspecific AND intraspecific positive
correlations between beak size and food size.
 Observation:
• Massive drought in 1977.
• Question: does drought affect bird-beak size
via natural selection?
 Testing Darwin’s postulates
• Postulate 1: Is the Finch population variable?
 Testing Darwin’s postulates
• Postulate 1: Is the Finch population variable?
Is some of the variation heritable?
• What does this mean?
 Main measure of genetic variation in a
quantitative trait…
• HERITABILITY: proportion of all variation due
to genetic differences among individuals.
(broad sense)
VP = VG + VE

Phenotypic = Genetic + Environmental

Variance Variance Variance

• Heritability = VG / VP = VG / (VG + VE)

Heritability

More often we want to measure the proportion of VP that results

only from VA.This allows us to predict phenotype of offspring
from the phenotypes of parents.

Narrow-sense heritability
= proportion of VP that results from additive
genetic variance.
 Heritability is a proportion
• Varies from 0 – 1.
• What does a heritability of zero mean?
• What does a heritability of one mean?
Calculating Heritability
Parent - Offspring Regression

The slope of the regression line provides information about the magnitude
Of narrow sense heritability.

Regress mean offspring phenotype onto mean parental phenotype:

 Measuring heritability
• Mid-parent offspring regression slopes (standard
method)
– Measure ‘trait values’ for parents.
– Measure ‘trait values’ of offspring (when grown)
– Idea…
• If differences between parents is genetic… then this will
be reflected in offspring.
• If differences between parents is environmental, it won’t.
Heritability in ground finches is high (0.65).
nd
(Darwin’s 2 postulate)

Heritability is high in both 1976

and 1978.
Why is there a difference in
size between 1976 and 1978?
Drought began.

Figure 3.11
 Postulate 3: Do individuals vary in their
success at surviving and reproducing?
• Most certainly:
• Drought (1977)
killed off 84% of
the birds.
 Why?
• Food abundance
went down
drastically.
 What about the food that was left?
• Seeds increased in
both size and
hardness. (bigger
beaks?)
Tribulus cistoides
began to dominate
 Postulate 4: Are survival and
Reproduction nonrandom?
• Only large big-beaked
birds can eat T.
cistoides.
• As a result,
big-beaked birds
survived better.
 Conclusion:
• All four of Darwins postulates are supported.
– Beak size is variable.
– Beak size has a genetic component.
– Survivorship is unequal.
– Survivorship is nonrandom.

Ergo… The population evolved by natural selection.

 Extra evidence

• IF the population evolved,

then the post-drought
birds should pass on their
big-beaked genes to their
offspring.
• Evolutionary change is
measured as the
phenotypic difference
between 1976 and 1978.
Evolution has been occurring over time
(note: not directional)
G. magnirostris
Large ground finch
specializes on
Tribulus cistoides
Some common misconceptions about
Natural Selection
 Important Points About Natural Selection

• 1) Adaptation happens over many generations.

 Important Points About Natural Selection

• 1) Adaptation happens over many generations.

– If an individual’s phenotype changes during its lifetime,
that is not adaptation.
 Important Points About Natural Selection

• 1) Adaptation happens over many generations.

– If an individual’s phenotype changes during its lifetime,
that is not adaptation.
– INDIVIDUALS DO NOT ADAPT.
– What adapts? (Remember definition of evolution).
 Important Points About Natural Selection

• 1) Adaptation happens over many generations.

– If an individual’s phenotype changes during its lifetime,
that is not adaptation.
– INDIVIDUALS DO NOT ADAPT.
– What adapts? (Remember definition of evolution).
• populations
 Important Points About Natural Selection

• 1) Adaptation happens over many generations.

– If an individual’s phenotype changes during its lifetime,
that is not adaptation.
– INDIVIDUALS DO NOT ADAPT.
– What adapts? (Remember definition of evolution).
• populations
**Natural Selection acts on individuals, but its
consequences occur in populations.**
 Cracking large seeds didn’t make beaks
grow larger. Small-beaked birds simply died.
2: Natural selection acts on phenotypes,
but evolution is simply changes in allele
frequencies.
No heritability? No
evolution.
3. Natural selection has no goal / it isn’t
forward-looking.
• Evolution occurs b/c of
genetic variation.
• Variation arises b/c of
RANDOM mutation.
• Selection cannot look
into the future and plan.
- Natural Selection Does Not Necessarily
Increase Complexity (it often does, but not
necessarily). BECAUSE IT HAS NO PLAN
4. While selection acts on existing
traits, new traits CAN evolve.
- i.e. corn oil content (after 100+ generations)
is over 3X the content of original parents.
4. While selection acts on existing
traits, new traits CAN evolve.
- New functions can arise.
- i.e. carnivorous plants: Butterworts: use
sticky trichomes to capture insects.
Trichomes are
known defensive
mechanisms for
herbivory
 Same is true for butterworts
- when hairs are removed in butterworts,
herbivory increases 18 times in the field.
 While selection acts on existing
traits, new traits CAN evolve.
- New functions can arise.
- i.e. carnivorous plants: Butterworts: use
sticky trichomes to capture insects.
- This is an example of an EXAPTATION: a trait
has evolved a novel use that didn’t exist in
the ancestor.
SECONDARY ADAPTATIONS can also arise
eventually making brand new structures:
secondary adaptations: additional modifications
that arise after an exaptation arises.
5. Natural selection does not lead to perfection.
Species are not “perfectly adapted.”
5. Natural selection does not lead to perfection.
Species are not “perfectly adapted.”
• Adaptations arise only if variation exists.
• Organisms may face contradictory patterns of selection.
5. Natural selection does not lead to perfection.
Species are not “perfectly adapted.”
• Adaptations arise only if variation exists.
• Organisms may face contradictory patterns of selection.
– Females MUCH prefer males with longer gonopodia.
5. Natural selection does not lead to perfection.
Species are not “perfectly adapted.”
• Adaptations arise only if variation exists.
• Organisms may face contradictory patterns of selection.
– Females MUCH prefer males with longer gonopodia.
– But, a long gonopod is “literally a drag”
5. Natural selection does not lead to perfection.
Species are not “perfectly adapted.”
• Adaptations arise only if variation exists.
• Organisms may face contradictory patterns of selection.
• Natural selection shapes organisms by culling from
available variants. There are developmental
constraints.

Developmental constraints are restrictions on the development of a particular

phenotypic trait. These are physical, mechanical or structural limitations as well as
irreversible commitments at key developmental stages that will limit or bias the
phenotypes that selection can work with.
“Natural selection is not a master engineer”

“It doesn’t produce the absolute perfection

achievable by a designer starting from scratch,
but merely the best it can do with what it has
to work with.” - Coyne
(Developmental Constraints)
Flatfish metamorphosis
 Example of imperfect design
• (Developmental constraints)
• “A conscientious designer might
have given [sea] turtles an extra
pair of limbs, with retractable
shovel-like appendages, but
turtles, like all reptiles, are stuck
with a developmental plan that
limits their limbs to four.”
• “No intelligent designer would
have given us [men] this tortuous
testicular journey… Our
testicular descent evolved later
as a clumsy add-on.”
6. Natural selection does NOT occur
to benefit a population / species.
- Self-sacrifice / altruism do not occur in nature. Every time a
seeming altruistic behavior is identified, a benefit to the
individual is found.

- (Chapter 12: group selection / evolution of social behavior)

6. Natural selection does NOT occur
to benefit a population / species.
- Natural selection occurs because SOME INDIVIDUALS (Not
species / populations) survive / reproduce better than
others.
6. Natural selection does NOT occur
to benefit a population / species.
- Natural selection occurs because SOME INDIVIDUALS
(Not species / populations) survive / reproduce better
than others.
- POINT: What is a single organism’s single biggest
competitor for resources / space / mates / etc.?
6. Natural selection does NOT occur
to benefit a population / species.
- Natural selection occurs because SOME INDIVIDUALS
(Not species / populations) survive / reproduce better
than others.
- POINT: What is a single organism’s single biggest
competitor for resources / space / mates / etc.?Other
individuals of the same population.
- Traits evolve regardless of their effect on the population /
species.
- Whatever effect they have on the species / population is a
byproduct.
 Sexual Selection
bad for the species?
 Finally, Don’t make this
mistake.
• No extant species is considered more
“advanced” / “higher” than another
living species.
• All life on earth arose from a single
origin… so all species have been
evolving for the same amount of time
(at least 3.5 billion years ago).
How Evolution Does / Does Not Work
• No modern species is the ancestor of another
modern species.
• (some sorta-exceptions)
 How Evolution Does / Does Not Work
• No modern species is the ancestor of another
modern species.
• Modern species are related…
• However, now on largely
separate evolutionary
trajectories.
 Phylogeny:
• evolutionary history of species / history of
speciation.
• Diagrammed as
a tree.
(using extensively in
this course)
 Words of warning
• be careful of the word "individual" and never imply that an individual can adapt.
• when discussing fitness and adaptation, refer to the environment within which a
trait has high (or low) fitness, or is an adaptation -- don't just refer to the trait.
• be careful of the words "population" and "species" and never imply that traits have
evolved for the good of a population or a species.
• never say or imply that any modern species is the ancestor to any other modern
species.
• never say or imply that there is a goal to evolution.
• never say or imply that a more complex trait must be more recently evolved.
• do not refer to "higher" and "lower" or "more advanced" and "less advanced"
modern species.
• do not assume, say, or imply, that a species or population must be perfectly
(optimally) adapted to its environment.