Studying Adaptation: Evolutionary adaptive significance requires

Analysis of Form and Function considerable ingenuity and effort.

Nectar Guide as Adaptive Form
- No hypothesis for the adaptive
- Consider a variety of methods
A trait, or integrated suite of traits, that
increases the fitness of its possessor is
called an adaptation and is said to be
adaptive.
Still other traits, or trait values, may
not be adaptive at all.
Analysis of Adaptation
value of a trait should be
accepted simply because it is
plausible and charming (Gould
and Lewontin 1979). Instead, all
hypotheses should be tested.
evolutionary biologists use to test
hypotheses about adaptations,
including experiments,
observational studies, and the
comparative method.
All Hypotheses Must Be Tested

The adaptive significance of some traits

may seem obvious
- Eyes are manifestly devices for
detecting objects at a distance by Buphagus erythrorhynchus and
gathering and analyzing light; in Aepyceros melampus
many animal species, individuals
with good eyesight will be better
able to find food and avoid
predators than individuals with poor
eyesight.

- Other traits offer more subtle

advantages. Understanding their
Red-billed ox-peckers have no effect on
the tick loads of cattle
Red-Billed oxpeckers maintain open
wounds on their hosts
Red-billed ox-peckers remove their hosts’
earwax
The oxpecker example demonstrates that
we cannot uncritically accept a
hypothesis about the adaptive
significance of a behavior, or any other
trait, simply because it is plausible or
because everyone knows that it must be
true. Instead, we must subject all
hypotheses to rigorous testing.
Other considerations to keep in mind
when studying adaptations are these:
- Differences among populations or
species are not always adaptive.
- Not every trait of an organism, or
every use of a trait by an organism,
is an adaptation.
- Not every adaptation is perfect.
Three methods to Test Hypotheses about
Adaptive Significance of Traits
- First: concern experiments
- Second: observational studies
- Third: explores comparative
method
Experiments
- Experiments are among the most
powerful tools in science. A well-
designed experiment allows us the
effect that a single, well-defined
factor has on the phenomenon in
question.
What is the function of the Wing Markings
and Wing-Waving Display of the Tephritid
Fly Zonosemata?
- The tephritid fly Zonosemata
vittigera has dark bands on its
wings. When disturbed, the fly holds
its wings perpendicular to its body
and waves them up and down.
- Entomologists had noticed that this
display seems to mimic the leg-
waving, territorial threat display of
jumping spiders (species in the
family Salticidae).
- Entomologists suggested that,
because jumping spiders are fast
and have a nasty bite, a fly
mimicking a jumping spider might
be avoided by other predators.
 - Because jumping spiders are
Zonosemata’s major predators
Greene and colleagues proposed
that the fly uses its wing-waving
display to intimidate the jumping
spiders themselves. The fly, in other
words, is a sheep in wolf’s clothing.
To run the experiment, Greene and
Mimicry of a predators behavior by
coworkers had to measure the responses
its own prey had never before
of jumping spiders and other predators to
been recorded.
the five types of experimental flies.
- Both mimicry explanations are
plausible hypotheses about the - When confronted with a test fly,
adaptive value of the fly’s wing- would the predators retreat, stalk,
waving display, but unless we test and attack, or kill?
them they are just good stories. - The researchers starved 20 jumping
Can these hypotheses be tested spiders from 11 different specie for
rigorously? two days. Then they presented one
of each of the five experimental fly
The researchers’ next step was to list
types to each spider, in random
alternative explanations for the behavior.
order.
Good experiments test as many
competing hypotheses as possible:
Hypothesis 1: the flies do not mimic
jumping spiders.
Hypothesis 2: the flies mimic jumping
spiders, but the flies behave like spiders to
deter other, non-spider predators. The Greene study illustrates important
points about experimental design:
Hypothesis 3: The flies mimic jumping
spiders, and this mimicry functions - Defining and testing effective
specifically to deter predation by the control groups is critical.
jumping spiders themselves. - All of the treatments (control and
experimentals) must be handled
- Biologists found they could cut the
exactly alike.
wings off a Zonosemata fly and
- Randomization is a key technique
glue them back on with household
for equalizing other, miscellaneous
glue. And they could cut wings off
effects among control and
a Zonosemata fly and replace
experimental groups.
them with the wings of a housefly
- Repeating the test on many
(Musca domestica), which are
individuals is essential. It is almost
clear and unmarked.
universally true in experimental
(and observational) work that
larger sample sizes are better.
Replicated experiments or observations
do two things:
- They reduce the amount of
distortion in the estimate caused by
unusual individuals or
circumstances.
- Replicated experiments allow
researchers to understand how
precise their estimate is by
measuring the amount of variation
in the data.
Observational Studies
- It is hard to imagine, for example,
how we could do a controlled
experiment to test alternative
hypotheses about why giraffes
have long necks.
- To do so, we would have to be
able to make giraffes that are
identical in all respects except the
lengths of their necks.
Behavioral Thermoregulation
Do nematodes show behavioral
thermoregulation in nature like they do in
the lab?
- The question is hard to answer for
an organism as small as C. elegans,
but more tractable for larger
creatures like lizards and snakes.
- To demonstrate behavioral
thermoregulation in nature, we
must show
(1) that the animal in question is
choosing particular temperatures
more often than it would encounter
those temperatures if it simply moved
at random through its environment,
and
(2) that its choice of temperatures is
adaptive.
Do Garter Snakes make adaptive choices
when looking for a nighttime retreat?
- Detailed study of the thermos
regulatory behavior of the garter
snake (Thamnophis elegans) at
Eagle Lake, California, Garter
snakes are affected by
temperature in same way as
nematodes.
- Garter snakes in the lab prefer to
stay at temperatures between 280C
and 320C.
 Body Temperatures of Garter Snakes in
Nature
- The main data for the study are the
lowest and highest critical
temperatures, called CTmin and
CTmax, that snakes can endure
briefly and survive.
- Compared the relative merits of
each of these thermoregulatory
strategies by monitoring the
environmental temperature under
rocks of various sizes, and at various
depths in a burrow, and by
monitoring the temperature of a
model snake left on the surface in
the sun or shade.
- By determining the options
available to snakes, and measuring
the frequency of each option in
the environment, the researchers
were able to show that the snakes
they observed were not simply
picking their retreats at random,
but were instead making an
adaptive choice.
The Comparative Method
Why do some bats have bigger testes
than others?
- David Hosken (1998) hypothesized
that large testes are an adaptation
for sperm condition. Sperm
competition occurs when a female
mates with two or more males
during a single estrus cycle, and
the sperm from the different males
are in a race to the egg.
- When formulating and testing
hypotheses about adaptation,
biologists must keep in mind that
organisms, and the lives they live,
are complex.
o Phenotypic plasticity and
o Trade-offs and constraints on
adaptation.
Phenotypic Plasticity
- Individual’s phenotype influenced
by its environment is to say that its
phenotype is plastic.
- When phenotypes are plastic,
individuals with the same genotype
may have different phenotypes if
they live in different environments.
- Phenotypic plasticity is itself a trait
that can evolve, and it may or may
not be adaptive. As with the other
traits we have discussed, to
demonstrate that an example of
phenotypic plasticity is adaptive,
we must first determine its function,
then show that individuals who
have it achieve higher fitness than
individuals who lack it.
Phenotypic Plasticity in the Behavior of
water Fleas
- Daphnia magna is a tiny filter-
feeding crustacean that lives in
freshwater lakes.
- Conveniently for evolutionary
biologists, Daphnia clone
themselves. This makes them ideal
for studies of phenotypic plasticity
 Trade-offs and Constraints
- Bat that have large testes help bats
win at sperm competition but
appear to impose metabolic costs
that lead to the evolution of smaller
and less energetically demanding
Phenotypic Plasticity in D. magna’s brains.
phototactic behavior - In an earlier passage we lamented
the male mosquitofish whose large
- An individual is positively gonopodium entices mates but
phototactic if it swims toward light slows his escape from predators.
and negatively phototactic if it
swims away from light. De Meester
measured the phototactic
behavior typical of different
genotypes of D. magna.
- The index can range in value from
-1 to +1. Female Flower Size in a Begonia: A trade-
- A value of -1 means that all the off
Daphnia in the test swam to the
bottom of the column, away from
the light.
- A value of +1 means that all the
Daphnia in the test swam to the
top of the column, toward the light.
An intermediate value indicates a
mixed result.

Monoecious- that is, there are separate

male and female flowers on the same
plant. The Flowers are pollinated by bees.
- The female flowers resemble the
male flowers in color, shape, and
size. This resemblance is presumably
adaptive.

Phenotypic Plasticity is widespread, and

perhaps underappreciated as an
adaptation.
- As Theodosius Dobzhansky pointed
out in 1937, “Selection deals not
with the genotype as such, but with
its dynamic properties, its reaction
norm, which is the sole criterion of
fitness in the struggle for existence.”
- Given that bees avoid female
flowers in favor of male flowers, the
rate at which female flowers are
visited should depend on how
closely they mimic male flowers.
- The ability to attract pollinators
should, in turn, influence fitness
through female function, because
seed set is limited by pollen
availability.

- Selection by bees favors larger

flowers, yet female flowers are no
bigger than those of males. Why
are female flowers not huge?