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BIO1130-Midterm 2 Notes

Introduction to Organismal Biology (University of Ottawa)

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4.2 Mendelian Inheritance

- Variation of genetic traits is explained by the different shapes genes (structural differences in
genome → variability of genetic traits) can have.

- Each gene occupies a specific locus (specific geographic locality on the chromosome) on a
given chromosome.
- DNA sequence of the locus can show variants​ (​allele​s​). So a gene can have different
forms. An individual will only have 2 alleles for one gene (hereditary factor) because they
only have 2 copies of chromosomes.

- All organisms inherit 2 copies of a gene (different or identical; one from father and one from
mother).

- A phenotype is the physical expression of a particular gene.

- ​The Law of uniformity​ of hybrids states that if both alleles of a locus are different, one of them,
the dominant allele determines the appearance of the organism whereas the recessive allele
has no notable effect on their appearance.

- ​The Law of segregation​ (where meiosis becomes important; believed to have evolved only
once) states that 2 alleles for a heritable character segregate during gamete formation and
randomly unite during fertilization (punnett squares show the probabilities of the possible
phenotypes).

- Testcross is used to identity the unknown genotype.

- ​The Law of independent assortment​ s​ tates that every pair of allele seperates independently
from the other pairs when the gametes are formed.

- If 2 traits are situated on the chromosome and the locus are really close to one another then
there is a high probability that they may be linked.

- The discovery of Mendel’s work had, initially, a negative impact on the theory of natural
selection. People preferred the discrete or discontinuous aspects of the traits associated with
Mendel’s ratios and rejected the continued variation observed in nature.

- Mendel dealt with transmission of traits from parents to progeny whereas Darwin dealt
with the evolution of populations.

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4.3 Initial Impacts of Mendel’s Theory

- Hugo de Vries formulated a “​Theory of mutations”​ which indicated that new species can
be formed in one or multiple steps through mutations that would cause major substantial
morphological modifications. Seemed to contradict Darwin’s theory of evolution.
According to de Vries:
- While working in a large field of yellow primrose, all of a sudden for no reason
whatsoever, a flower came out red. Hugo said “aha mutatation → aha new
species → therefore Darwin is wrong because we do not need variation just a
mutation to create a new species. The problem with this is that even if you have a
mutation that produces a different morphology, you need to see if that morph will
actually reproduce with the yellow primrose or if it needs to reproduce with
another red one. It was later understood, that the red flower was no more than a
chromosomal defect in the flower.
- Discovered that most mutations were harmful or neutral and that only a few seemed to
be beneficial in an evolutionary sense, specifically mutations that were favoured in
certain environments.

4.4 Neo-Darwinism and Modern Synthesis

- We were finally able to quantify natural selection and evolution.

4.5 Modern Additions to Mendel’s Laws

- Incomplete dominance​ (one allele is slightly dominant over another allele) → A

combination of the phenotypes of each allele is expressed (i.e chestnut horse + cream
horse → Palomino horse)
- Codominance​, where both alleles are dominant and are equally expressed (i.e black
chicken + white chicken → checkered chicken)
- Polygenism​: When a phenotypic trait is under the combined action of two or multiple
genes (very common → i.e skin colour, height)
- Common in continuous traits (traits where there does not seem to be a discrete
value to it
- Epistasis​: When the effect of a gene at a certain state (i.e recessive homozygous) hides
or blocks the expression of another gene.
- Pleiotropy​: When a gene influences more than one trait.
- Certain pleiotropic gees are lethal when homozygous:
- Manx cat: The Manx gene (M) shows an incomplete dominance to the normal
gene (m) (with tail)
- The gene responsible for the lack of a tail is lethal (embryo reabsorbed) when
homozygous.
- The heterozygous cat do not have tails or have short tails.

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M m

M MM* lethal Mm

m Mm mm
* never gets to be born → the whole vertebral column does not develop properly and therefore
embryo is reabsorbed

Chapter 5: The Evolution of Populations

5.1 Hardy Weinberg’s Principle

- Weinberg and Hardy demonstrated that, under certain conditions, allele frequencies in a
population stay constant from one generation to the next. Thus, no evolution. (null
hypothesis of evolution).
- A principle that describes a hypothetical population that does not evolve
- Conditions of a population in Hardy-Weinberg’s equilibrium:
- There are no mutations
- Mating is done randomly
- The size of the population is extremely large
- There is no genetic (gene) flow (no migration of alleles between populations).
- There is no natural selection

5.2 Mutation

- Changes in the DNA sequence of an organism.

- This is the source of genetic variability​. Mutations are:
- Random; do not arise as a need but may be favoured by natural selection.
- Transmissible (only if they are in the sex cells/gametes).
- Frequent throughout the gene pool but rare at each locus.
- Will influence allele frequencies but is a weak evolutionary force from generation
to generation, especially in large population because they are rare at each locus.
- Point mutations (addition, deletion or substitution of a base)
- Rates of mutations vary greatly according to groups of organisms or genes (bacteria has
a high rate of mutation)
- Measured in pair of bases per generation (pbg)
- Even if a small portion of these mutations is advantageous, it offers a good
genetic basis for evolution over millions of years.

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5.3 Assortative Mating

- Random mating: panmixia (maintains Hardy-Weinberg Equilibrium)

- Also known as a panmictic population:
- American eel are panmictic populations. After a few years, these eels will actually
do a migration toward one specific area → Sargasso Sea (the site of
convergence of all the populations and random mating would occur). Looking at
the population at each river, there doesn’t seem to be any kind of genetic pattern
(unlike salmon which migrate back to their place of origin, eels do not migrate
back to the same original location → therefore random mating occuring in the
Sargasso Sea)
- Assortative Mating: Choice of partners in relation to the phenotype (modifies H.W.
equilibrium):
- Positive AM: more frequent mating between similar individuals than expected by
chance. (Effect → increases homozygosity)
- Autogamy or selfing of plants (pure lineage of Mendel)
- Geographical proximity of individuals (e.g. population of mice in a barn)
- In humans: mating according to height, skin colour, religion, etc.
- Negative AM: more frequent mating between individuals that do not look alike
than expected by chance (Effect → increases heterozygosity)
- Many mechanisms in nature that promote heterozygosity because there is
an advantage because then you can hide some genes in the
heterozygote that may be useful afterwards.
- Most flowers cannot self reproduce and hence use pollination to mate.
Some flowers develop immunological or biochemical ways to reject their
own pollen in order to prevent self fertilization.
- Assortative mating does not change allele frequencies; it changes genotypic
frequencies
- When paired with natural selection, positive assortative mating can lead to an
increase in homozygosity and a loss of genetic variability:
- 1. Phenomena of endogamic depression (black guppies) since the
harmful alleles will express themselves.
- 2. Natural selection will purge the population of a portion of its harmful
alleles
- Thus, a loss of genetic variability compared to panmixia

5.4 Gene Flow

- populations are group of individuals that usually live in specific geographical areas.
- The concept by which individual from one population can get into another population
(almost equivalent of a mutation → addition of new alleles/genes into the receiving
population)

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- Therefore it will have an effect on the genetic makeup of the recipient population
as genotypic and allele frequencies are changing = evolution
- Important factor (migration of individuals) to investigate when looking at the
evolutionary history of a population
- You cannot have gene flow between species (i.e ostrich → geese) only between
populations
- What is the ultimate consequence of migration?
- From a genetic perspective, the ultimate consequence of migration is to add new
alleles to a population.
- If migration is done between populations over a long period of time, both
populations will end up with the same genetic variability. Gene flow has a
tendency to standardize (the genetic pool of the populations involved)
populations to have the same genotypic and allele frequencies
- A powerful mechanism that is important in conservation biology
- Example of Gene Flow:
- There are 2 populations of Great tit on an island in the Netherlands: Central
Population and Eastern Population. And it seems that the central population is
the main recipient of migrants from the mainland and because of that, the
population of Great tit in the centre of the island has a lower reproductive
success than the eastern population, which is a well established population on
the island. Individuals from the mainland are not well adapted to an insular life
(life on an island). Therefore, the number of females that survive and eventually
breed is much lower in the central population than the eastern population.

5.5 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.
- Chance will have increasingly more impact on a population as the size of the population
gets smaller
- In ​small​ populations, genetic drift will trigger a decrease in genetic variability and
a decrease in heterozygosity.
- In a ​large​ population, genetic drift will cause little changes to the allelic or
genotypic frequency of a population.
- Thus, the larger the sample, the smaller the differences will be between expected
frequencies and observed frequencies
- If there are no other processes (mutation, migration, or selection) that will affect allelic
frequencies at a particular locus, genetic drift will eventually result in the ​fixation​ of an
allele and the elimination of all others for this locus.
- If no other evolutionary processes are acting on allelic or genotypic frequencies, then the
probability that an allele will become fixed is ​equal to its frequency.

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- A population bottleneck is when a population gets reduced to a few individuals who bear
only a portion of the genetic variability that was found in the original population. Thus,
increasing the risk of extinction.
- Example: Great prairie chicken
- Large prior population had a high genetic variability of 5.2 alleles per
locus and as a result had 99% of eggs hatched. After a population
bottleneck, the remaining individuals had a lower genetic variability of 3.7
alleles per locus and consequently had <50% of eggs that hatched. This
is because when you reduce genetic variability, homozygosity increases
which means that a lot of individuals that have the same genotype
reproduce together → recessive alleles will become homozygous and
some of the recessive alleles have negative impact on survival and
because of its prevalence within the population, affects the whole
population’s adaptability.

- Example of genetic drift: ​Founder effect

- When a few of individuals from a larger population create a new colony in a
different habitat, the genetic makeup of the colony differs from the source
population. If there are few negative alleles present in the population that
migrated to establish the new colony, because the small population will
interbreed, then there is a higher probability that some diseases may become
more prevalent.

5.6 Natural Selection

- A process by which the individuals with certain hereditary particularities survive and
reproduce in larger numbers than other individuals (differential reproductive success)
- Alleles favoured by selection are more abundant in offspring than in parental generation
- The action of natural selection on the frequency of an allele can be cancelled by the
action of mutation, genetic drift, and migration.
- The only evolutionary mechanism that aids the survival and the reproduction of
organisms in their environment
- ONLY hereditary variations constitute the basis of natural selection
- Example of non-hereditary variations: Map butterflies
- This species has varying colourations depending on whether it is summer
or spring and these differences in colouration are not genetic. They are
ENVIRONMENTAL. Colourations are due to the difference in the plants
they will feed on (they feed on different plants in the summer and spring).
Hence, natural selection cannot act on this because the colouration is
characteristic linked with the environment (no genetic basis).

- Variability in yarrow height according to altitude (Genetic differences or environmental

effect?):

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- At lower altitudes, they are very tall but are shorter at higher altitudes. To test
this, you take the identified seeds (from both altitudes) and put them in a
greenhouse and offer them the same environmental conditions (same
temperature, same nutrients, same daylight exposure) and you observe their
growth. If the plants grow to the same height, then you know that the difference in
height observed at different altitudes are only based on environment

- The ​Adaptive value​ (selective value, fitness) of a genotype corresponds to the

contribution of an individual to the genetic pool of the next generation when compared
with the contribution of other individuals.
- Types of hereditary traits that natural selection can act on:
- Qualitative (discrete variation: colour)
- Quantitative (continuous variation: height, weight)
- Polymorph​: populations showing distinct morphological types of genetic variability

- If we take the case of a continuous trait, then we usually divide natural selection into 3
types:
- ​Directional selection​: the average proportion of individuals with a specific trait
increases (bell curve shifts toward the favoured trait)
- Examples: Medium Ground Finch beak size; random point mutations in
some mosquitos that protect it against DDT (develop resistance and pass
this trait/allele to offspring → population becomes resistant)
- Disruptive selection​: there is selection against middle values and selection for
extreme values (mean value of population will not change; if disruptive selection
is really really strong and certain conditions are met, it may lead to speciation)
- Beak size of Pyrenestes ponceau: selection/drought against intermediate
beak sizes and favoured wide or narrow beak sizes (either because beak
could not handle intermediate-sized seeds or drought caused no
intermediate-sized seeds to occur).
- Stabilizing selection​: selection against extreme values in the population.
- In the 1930s and 1940s (in England), the ideal weight for newborns was
about 8 pounds (3.6 kg). That meant, babies at this weight had a higher
rate of survival than babies of a larger or smaller weight. Today this
stabilizing selection has almost disappeared in wealthiest countries
(medical advances).
- Another example: all mammals have 7 cervical vertebrae (in neck).
People with more or less will have a pleiotropic effect → people born with
this is usually associated with diseases and survival rate is low

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5.7 Preservation of Genetic Variability in Nature

- The above 5 topics are evolutionary mechanisms that can modify the allelic and
genotypic frequencies of a population. All are motors of evolution, but only natural
selection will create adaptation
- Genetic variability is maintained in nature (​polymorphism​) by the following:

- Diploidy:
- A considerable portion of the genetic variability of diploid individuals is hidden
from natural selection:
- The mass of little expressed or unexpressed genes (in phenotype of
organism) in heterozygotes is called the ​genetic load
- It is the cost associated with maintaining and storing genetic
variation.

- Balanced Polymorphism:
- The heterozygote advantage​:
- When heterozygous individuals (have an advantage) have more offspring
than homozygous individuals.
- Example: Sickle cell anemia (incomplete dominance);
- S dominant; s recessive
- Homozygous ss = sickness; high mortality before the
reproductive age.

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- The allele frequency “s” is particularly high in regions

where cases of malaria are high
- The heterozygote Ss is more resistant to malaria than SS

- Frequency-dependent selection:
- The selection is dependent on the inverse frequency meaning that
sometimes there is advantage to being rare (rarest phenotype is favoured
by selection).
- Example: type of fish that is found in some African freshwater
lakes that actually feed on the scales of other fish. Within this
population of fish, you will have fish that have mouth that is curved
to the left that will attack the right side of the fish and there are
other fish with a mouth twisted to the right which attacks the left
side. If one side of the prey gets attacked often, the prey will
become more watchful of fish coming from that side than the other
side. This will increase the success of fish that attack the opposite
side thus leading to a population with a greater proportion of fish
that attack that side and vice versa (cycle continues).
- The selection that depends on the positive frequency. The phenotype that
is more abundant is always favoured (opposite of previous type).

- Neutral Variation
- A good portion of the genetic variability found in genes do not show selective
advantages or is not affected by natural selection (e.g pseudogenes).

- External mechanism (or ecological):

- The result of simultaneous impacts of different selective pressures. Population
are submitted to all kinds of selective pressures.
- Temporal changes in selective pressure (changes through time i.e climate
change)
- Habitat mosaic → not all individuals of a species live exactly in the same habitat
(some species are very widespread)
- Non-assortative mating

5.8 Sexual Selection

- Initial formulation of the concept: Charles Darwin

- Sexual dimorphism → difference between the males and females of a species
- A form of evolution in which individuals that possess some specific hereditary traits are
more susceptible than others to find partners. (i.e female peacocks choose mates that
have the most dots on their feathers)
- Intersexual​ selection (choice of sexual partners based on traits indicating the
quality of the genetic baggage/health of the other sex):

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- The case of the widowbird: Do the females have a preference on the

length of the tail features of males?
- Sexual dimorphism: female is brown/yellow/green with marble colour
whereas the male is black and most of them will have a very long tail.
Females stay on the nest and since they are savanna/marsh type of
habitat, their colouration provides camouflage. The males are territorial
and will try to attract females to their territory. To determine what a female
widowbird looks for in a mate, a test was established. 3 groups of males:
cut tail, no change, and attached tail of cut tail to normal. Males with
longer tail were chosen as it is seen as an indicator of good genetic
health.
- Intrasexual​ selection (selection/competition between individuals of the same sex)

Chapter 6: Adaptation

6.1 Adaptation

- Adaptations are the result of natural selection (the only process that creates adaptations)
- They are sometimes complex, have complex functions and a complex history (ex. May
be a result of evolutionary compromises; symbiosis; common ancestry)
- Why is our species the only Primates that can choke (food entering respiratory tract) on
food?
- An evolutionary compromise. A human’s morphology consists of a flat face, long
neck and a fairly short mouth as we feed on substances that are quite soft → in
fact jaws got smaller because for the last 2 million years because we have been
eating meat and for the last million years we have used fire to tenderize the meat
and we used tools to cut the meat which makes our chewing process easier and
therefore our lower jaw has evolved to become smaller. We also have teeth that
were once useful but now do not serve any significant function (i.e wisdom teeth
→ painful because jaw is getting smaller faster than the number of teeth we
have) and in terms of the nostril (we rely less on our sense of smell and more on
sight). If we look at the back of the throat, there is a space between the soft
palate and the epiglottis (this space is what distinguishes us from the chimps),
and this space creates the possibility that there will not be an automatic closing
action of the epiglottis, and a false nerve reflex, food will enter the trachea. This
is long vertical tube is also the reason for our ability of articulated language that is
so unique to us. Despite our vulnerability to choking, the advantage gained from
speaking has a higher adaptive value or compensates for this.

- Burying beetles → will find corpses somewhere (i.e mouse) and first bury it by softening
the soil around it and cover it with saliva to prevent it from smelling and attracting other

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competitors. When this is done, they will lay a few eggs by the corpses and the eggs will
hatch and the decomposing corpses will serve as a food source (parent will feed the
larvae). When the larvae gets large enough, the larvae will actually migrate inside the
corpses and complete their cycle/transition into a beetle.
- Transport mites → seem to be using the beetle as a means of transportation in order to
take advantage of flight to access a new food source as the mites also do their life cycle
on the corpses.
- Symbiosis: interaction between 2 species and probably an indication of coevolution
(adapted to one another)
- The case of the burying beetle and transport mites is ​commensalism​ (only one species
benefits while the other is neither helped nor harmed). New science hypotheses that it
might instead be a case of ​mutualism​ (both partners benefit) where the benefit for the
beetle is that the mite may in fact get rid of some of the competitors of the beetle on the
corpses meaning that the mite is selective toward the larvae of the beetle and would
help keep the food source for the larvae of the beetle by eliminating all other competing
larvae on the corpses.
- Another example of commensalism is that the fact that we have on our body in every
square centimeter of the surface of our skin, billions of bacteria that are present and
actually feeding on our sweat and dead cells.

- Parasitism​: one organism (the parasite) gains, while the other (the host) suffers.
- Flatworm parasite (in sushi): you don’t feel anything until it becomes quite large;
found in flesh of salmon; they have 3 hosts which are necessary for the parasite’s
life cycle: salmon, in other species of fish, and zooplankton. Infection by the
flatworm results in sickness, deficiencies in some vitamins.

6.2 Studying Adaptations

- We may think of organisms as perfectly tailored to their environment but all traits of a
living being are not necessarily adaptations to the current environment: ​concept of
exaptation ​(feature that has not necessarily evolved in recent species but might have
evolved in the past).
- Adaptation​: a feature produced by natural selection for its current function.
- Exaptation​: a feature that performs a function but that was not produced by natural
selection for its current use. Perhaps the feature was produced by natural selection for a
function other than the one it currently performs and was then co-opted for its current
function. For example, feathers might have originally arisen in the context of selection for
insulation, and only later were they co-opted for flight. In this case, the general form of
feathers is an adaptation for insulation and an exaptation for flight.
- Methods aimed at corroborating or refuting adaptive hypothesis:
- Direct study of natural selection (eg. peppered moth, medium ground finch,
human adaptation to altitude)

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- Comparative approach: indicator that traits have evolved independently or are

the result of a common ancestor. (e.g. carnivorous jaws vs herbivores jaws)
- Morpho-functional study of a trait (e.g. evolution of fur or feathers)

- If someone tells you that the lion’s jaw is beautifully adapted to its way of life, this is
wrong because the lion did not evolve this jaw, it was evolved in a common ancestor.
This is NOT an example of exaptation because the function of the jaw has stayed the
same (cutting meat). In a carnivore, articulation connecting the lower jaw to the skull is ​at
the same level​ as the row of teeth (solid articulation).

- Herbivores have this trait of the jaw where all the teeth touch each other at the same
time and the jaw joint is very loose allowing for easy movement of the jaw. This
adaptation is evolved from a common ancestor of all herbivores.

- Dinosaurs started very small and ended very large. The largest species of dinosaurs
were found in the last 10-15 million years of their history. The T-rex evolved from another
species and in fact evolved from a fairly small species (same morphology but different
sizes). Therefore to say that the morphological characteristics of the T-rex evolved in the
T-rex is ludicrous (its features/adaptations were evolved from smaller common
ancestor). To understand why T-rex had small arms → need to look at common
ancestor. Since there was no selection for or against it, the T-rex kept the small arms.
The reason for the T-rex’s size can be explained through the arms race concept (i.e.
cheetah vs gazelle → arms race for speed).

- Example of exaptation (adaptation whose current function is not the one for which the
structure initially evolved): Fins and limbs
- The initial function of fins was to ​stabilize​ the fish body in its environment; it is an
adaptation.
- Fins are the ancestor of limbs; in terrestrial animals (tetrapods), the limbs (that
have evolved from fins) are used to support the body in an aerial environment.
The limbs of the tetrapods are thus an ​exaptation​ for terrestrial mobility
(​locomotion​). Fish however, did not need to support their body as the dense
aquatic environment supported their body for them.

- Hair evolution: Adaptation or Exaptation ?

- Hair is the main characteristic that unites mammals. It has an important role in
thermoregulation because it prevents loss of heat by creating this layer of warm
air.
- Case study: elephant
- Elephants have areas of their skin with sparse hair and with no hair at all
(no fur). Hot spots show that elephants do not lose heat in a homogenous
fashion. Why do elephants have hair in an hot environment where fur is
not necessary? Why didn’t they lose all their hair (i.e whales)? Research

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concluded that having sparse hair on the skin, is actually more efficient at
losing heat than having no hair at all. This is explained through an
engineering model/principle: pin theory where a sparsely hairy surface is
better at losing heat (25% more heat lost) than a flat surface. Hence,
maybe the density of hair and not hair itself, an adaptation to preserve
heat. But for what reason (or how) did hair evolve? → not to preserve
heat because one strand of hair does nothing to preserve heat → so it
had to have evolved for another function. Therefore, hair might have
actually evolved first to ​lose​ heat. Another initial function of hair might
have been associated with the nervous system. We know that in some
animals, there are whiskers that have nerve endings that allows the
animal to orientate itself in the dark and pick up stimuli from the
environment (so maybe hair evolved initially as a sensory structure). In all
probability, the statement that hair is important to us in terms of
preserving heat is an exaptation. In fact we can say that it is not the hair
itself but the fact that we have hair is why we preserve heat.
- Therefore, there are 2 possibilities for the original function of hair:
- 1. Dissipates heat
- 2. Sensory structure

- Feathers: Adaptation or Exaptation?

- We often associate feathers with the act of flying. But why did feathers evolve?
What was the initial function/adaptation? It did not evolve for flying, because the
feature evolved in animals that were large (therefore feathers did not evolve for
flight). The organization of feathers may have evolved for thermoregulation but
what did the feather itself evolve for? Initial adaptation could be for losing heat
but this does not work for dinosaurs → because they were cold blooded (could
not generate their own body heat, ectotherm). Another possibility could attributed
to sexual selection (appearance of feathers to attract mates). Could be a sensory
structure. To say that a structure evolved for the beautiful purpose we see now is
often not science based (need to know origin of function).

- Inner Ear: we have small bones that allow us to keep our equilibrium and are soaking in
some liquid that is very important for our health when this gets infected. Looking at the
history of these bones, we see that they are actually bones of the jaws of fish that have
been recycled (evolutionarily speaking) into a new function. So the ear ossicles help
hearing (exaptation) but the original adaptation is to catch food in fish.
6.3 Evolution and Developmental Genes

- Heterochrony​ : Changes in the speed or synchronization of the developmental phases.

- Some parts of the body grow faster or at a specific time. In our developmental process,
we have genes that express themselves and stop expressing themselves.

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- Example of paedomorphosis: Necturus

- It is a very interesting salamander. It does not have a terrestrial phase. The
larval/aquatic phase acquired the capacity to develop fully functional reproductive
organs. Essentially, the accelerated development of the reproductive organs
overtook the development of the body → this created an aquatic salamander
capable of reproducing. Adaptation can also be the result of the change in
development timing.

Chapter 7: Geographic Variation and Speciation

7.1 Inter-population Genetic Variation

- We observe geographic variation in all species.

- Example: One species, two populations of caribous
- 2 herds that overlap slightly in terms of their distribution yet the herds are slightly
different morphologically and in spite of their overlap there appears to be no
crossing between the 2 populations of caribou → ​positive assortative mating
(individuals of each population have a tendency to mate with members of their
own population)

7.2 Models of geographic variation

- Polymorphic​ species: species formed of several geographic groups which differ from
each other by traits that are easy to recognize (sometimes called morphs, forms or
subspecies). (i.e Greater racket-tailed drongo in south east Asia)

- Ecogeographic​ rules: models of geographical variation that follows climatic gradients for
many species within a group of vertebrates within a group of vertebrates
- Bergmann’s rule​: In endothermic animals*, the population living in northern
regions of the geographic distribution of the species will generally have a ​larger
size. The reason for this is debated among the scientific community; but the
accepted reason is thermoregulation. If you are really small, you have a large
surface area for a small mass (large surface area relative to mass) so in terms of
animal that produces heat, you are losing your heat much faster. Northern
animals have a much smaller surface area to mass ratio, which means they lose
heat at a much lower rate. (i.e white-tailed deer)
- 72% of birds and 65% of mammals follow Bergmann’s rule.

- Allen’s rule​: In birds and mammals, the northern population (or species*) will
generally have ​short and massive​ extremities, whereas the southern populations

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(or species) will have ​longer and thinner​ extremities. This is linked with
temperature dissipation and/or preservation of heat.

- Gloger’s rule​: within a species of endotherms, more heavily pigmented forms

tend to be found in more humid environments, e.g near the equator. An
explanation for this, in the case of birds, appears to be the increased resistance
of dark feather to feather/hair degrading bacteria. Feathers in humid
environments have a greater bacterial load, and humid environments are more
suitable for microbial growth; dark feathers or hair are more difficult to break
down.
- Therefore pigmentation of species is dependent on its habitat: darker →
humid/dense habitat, more bacteria that attacks bird so more melanin acts as a
defense (antibacterial) mechanism.

Evolution of the loss of fur and skin colour in humans

- An example of geographic variation in humans is differences in skin colour.
- Sahelanthropus tchadensis is the oldest hominid (bipedal posture) known. The main
feature that unites all hominid species (all human species) as one large group with a
common ancestor is their bipedal posture. First human species had pale/pink skin
covered with dark fur.
- All traits that will be discussed, except the evolution of pale skin, are traits that did not
evolve in our species (we inherited these characteristics).

→ ​Australopithecus afarensis ​(3.7-3 Myr)

- Lived mainly in tropical forests; bipedal and arboreal (spent a lot of time on trees shown
by long arm structure); diet of fruits, tubers and leaves; sedentary way of life (no capacity
to migrate)
- In the last 60 million years, the average temperature decreased. In the last 3 Myr, there
has been an ​important cooling​ of the Earth’s temperature. Associated with this change in
temperature is the fact that the area of Africa with the most human fossils has always
been an area of geographic instability (due to presence/contact between tectonic plates).
The cooler temperatures caused the tropical forests to recede and lead to the formation
of savannas and deserts and mountainous habitats. Some species of humans adapted
to the new, open habitat (this new habitat was a strong selective pressure to add
something to the diet → meat). The quest for water also became a problem and required
a capacity to move around easily (mobility)

→ ​Homo ergaster ​(= ​H. erectus​) (1.2 Myr)

- Less fruit available; meat added to the diet
- Walks longer distances for prey and water.
- Adapt to life on the plains
- More active lifestyle compared to sedentary species (​hunter-gatherer​)

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- Natural selection acts on the shape of the body/skeleton → leaner, longer limbs (ability
to run)
- Abundance of ​sweat glands and less fur​ to have a more efficient
thermoregulation
- Loss of fur increased efficiency in terms of evapotranspiration.
- In a furry animal, there is mostly apocrine glands which produce oily sweat and
as a result perspiration is difficult
- In humans, there is mostly ​eccrine (waterly) ​sweat → easy perspiration
- Carnivore thermoregulation: ​panting​ and eccrine sweat glands only on the ​pads​ of paws
or mouth (panting).
- Prior to humans losing their fur, the fur served as protection against UV rays and
protection against bacteria, and abrasion (these functions were then assumed by the
skin → nature of skin changed).

→ Selection in favour of darker skin (1.2 Ma) ​at the same time​ as fur loss.
Natural selection favoured the individuals having thicker, darker and more acidic skin (more
melanosomes = more melanins). Darker pigmentation offers antibacterial properties (acidic).

- Protection against: Ultraviolet rays (UV), drying (loss of water), bacterial attacks and
vitamin deficiencies.

- Another consequence of losing fur is the impact on communication. Animals use their fur
for communication → they display some emotions by the shape of their fur. By losing our
fur, we lost this capacity to communicate. Scientists think that the ability to communicate
was transferred from our fur to our capacity to speak and we kept hair in specific places
on our face that allow us to transfer our emotions via facial expressions. So
communication was another function previously associated with fur that has been
transferred to naked skin and maybe implies the evolution of sound as a means of
communication or even facial expressions.

- For over 1 Myr, the skin of ​all​ hominins species including ​Homo sapiens​ was dark.

- Hypothesis to explain evolution of skin colour:

- Reminder: We need to find an ​inherited​ trait that gave those who had it a positive
survival and reproduction differential​ (adaptation) from generation to generation
compared to other members of the population. (usually mortality associated with
any type of skin disease associated with UV radiation, you die at a fairly old age.
That means that you can your family before you actually die and thus this is not a
good mechanism to explain the evolution of skin colour → ​no effect on survival
and reproductive differential ​→ need something that affects someone early on in
their life → must affect reproductive success)

→ ​Folic acid and dark skin​:

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- Folic acid (vitamin B9) is destroyed in skin by excessive UV rays.

- vitamin B9 deficiency: serious and possibly lethal developmental malformations (spina
bifida), poor wound healing, disturbed immune system, sperm malformation.
- A vitamin deficiency at a young age and, particularly, in pregnant women would have a
direct impact on the ​survival and reproductive success of the individual​.
- Thus, we can say that ​dark skin​ offers a protection against UV rays that offers a survival
and reproductive advantage (​that’s an adaptation​)

→ ​Vitamin D3 and pale skin​:

- Vitamin D3 is synthesized in the skin by UV rays. It helps in the absorption of calcium in
the gut. (Deficiency: rickets, plus several other diseases,...).
- A deficiency in vitamin D3 would have a direct impact on the reproductive success of
affected individuals.
- Pale skin in low UV radiation would maximize the absorption of UV rays and the survival
of human population in areas of UV deficiency (​Adaptation​)

-------------------------------------------

- The great human migration that started ​100 000 years ago​ has seen humans invade
increasingly northern habitats more recently (especially in the last 40 000 years) and
evolve paler skin to maximize UV absorption in area where UV radiation is weak for
adequate ​synthesis of vitamin D​ by the skin.
- In areas with an important annual deficit of UV rays, colonization by humans (10 000 to
15 000 years ago) was made possible because humans compensated for deficiencies in
vitamin D3 through ​diet​ (hunting, fishing and domestication)

→ Conclusion:
- Loss of fur (and change in morphology) in hominins is linked to ​lifestyle​ changes due to
climate change, more than 1.2 Myr ago.
- The skin colour became ​dark​ quickly after the loss of fur. The skin remained dark for
more than 1 Myr.
- The evolution of pale skin in some human population is linked to the colonization of
habitats further north during the last 100 000 years. This paler skin is an adaptation to
maximize the absorption of UV rays in zones where radiation is low; this allowed the
healthy synthesis of ​vitamin D3​.
- Note: individuals with darker skin in regions of high UV intensity synthesize
Vitamin D3 at a much ​slower rate​ than individuals with pale skin.
- A selective advantage of darker skin in regions of high UV intensity is to minimize the
degradation of ​folic acid​ by UV rays.

- The colour of skin is:

- A variable ​polygenic​ trait which explains the variability of pigment intensity.
- Only an adaptation to ​UV radiation​.

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- Only an indicator of the ​environment​ in which populations have lived.

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