1Segment 1: Darwinism

The Early Case for Evolution

Charles Darwin was an English naturalist who began studying biology as a student in the 1820's.

- Darwin’s famous voyage on the HMS Beagle (1831-1836) greatly influenced his view of
biology and culminated in his theory of evolution through natural selection.

- at that time in England, the prevailing explanation for the origin of life was the Theory of
Special Creation

- this theory states that:

1) all organisms were created by God in the 6 days of the creation as described in Genesis of the
Old Testament

2) all species had remained unchanged (immutable) since the creation

ie: animals and plants remain unchanged in form from the moment of their initial appearance on
Earth (fixity of species).

3) the creation event was recent

eg: Archbishop James Ussher (1664) calculated the Earth to be 5668 years old and was created
at 9AM on October 26, 4004 BC.

- starting about this time in Europe, the idea of evolution began to supplant the theory of
creationism

Several types of data being collected during this period began to support the theory of
evolution:
1) Relatedness of Life Forms
Anatomists began to realize that vertebrate embryos were very similar (especially in early
development)

- Darwin noted in his The Descent of Man (1871) that humans are developed from an egg that is
no different from other vertebrates

- he included an illustration of a dog and a human at about the same stage in development to
illustrate that:

“The (human) embryo itself at very early period can hardly be distinguished from that of
other members of the vertebrate kingdom.”

- human embryos go through a fish-like stage with gill-like pharyngeal pouches, a tadpole-like
stage with a tail and a stage when the embryo has fur

- the study of embryonic development is called ontogeny

- a common expression among evolution biologists is:

“ontogeny recapitulates phylogeny”

- that is, the basic form of an organism can be observed in its earliest embryonic development;
modification to these basic forms occurs in later development

- therefore, early embryonic development reflects earlier evolutionary forms and later
development reflects modification to the basic form

again: early developing human embryos exhibit pharyngeal pouches that are homologous with
the gill slits of fish

eg: later developing embryos exhibit a dense coat of fur

Like animals, plants also undergo embryonic development and are studied in the same way and
for the same reasons that animals are

eg: lima beans are plant embryos surrounded by endosperm

eg: floral developmental studies are carried out using the scanning electron microscope

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Similarities that have a common origin are called homologies.

- vertebrate homologous structures develop from the same group of embryonic cells

- the same is true in other organisms such as plants

- to the question: Why should these organisms share the same developmental pathways?

- Darwin, in his The Origin of Species (1859) responded that descent from a common ancestor
was the most logical answer...

“What could be more curious than the hand of a man, formed for grasping, that of a mole for
digging, the leg of a horse, the paddle of the porpoise and the wing of the bat, should all be
constructed on the same pattern, and should include the same bones, in the same relative
positions.”

Structures that are similar but are not descended from a common ancestor are called analogies

- these structures are the result of a process called convergent evolution

eg: the wing of the bat and the wing of a fly

eg: there are 3 unrelated families of pitcher plants that are similarly shaped and supplement their
nutrition by capturing insects

- that is, when unrelated organisms solve a common problem and take on similar characteristics,
this is convergence

2) Change Through Time

- at this time, evidence began to mount that species were not immutable

- many paleontology studies documented the close relationship between extinct fossil species and
their living relatives from the same geographical area

- this pattern is known as The Law of Succession

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eg: Darwin analyzed the armadillos of Argentina and their relationship to the extinct
glyptodonts

- additional evidence that organisms are not immutable comes from vestigial structures

- these are structures that were once useful in ancient species but are not longer necessary fro the
present species

- however, not enough evolutionary time has lapsed for the structure to be completely eliminated

eg: pelvis and femur in extant whales

aside: whales evolved from wolf-like animals starting about 50 million years ago; fossil whales
from about 40 million years looked similar to modern whales except that they had forelimbs
rather than flippers

eg: tail bone in humans

eg: tissue bulbs in the eye sockets of blind salamanders

- these structures suggest that organisms are modified in time

eg: projecting points in some human ears as discussed in Darwin’s book The Descent of Man
published in 1871

- extrinsic muscles are responsible for the movement of ears in some animals (e.g. dogs, cats,
cows, etc.)

- these muscles direct the ear to enhance sound sensation

- in humans and apes, these muscles are in rudimentary state

- some people can move or “wiggle” their ears with these muscles while others can not

- the inability of humans to move their ears is partially compensated by their ability to move their
heads in the horizontal plane

- some people have a projecting point along the outer folded margin (helix) of the ear

- these points appear to be vestiges of the tips of formerly erect ears and represent reversals in the
individuals that possess them

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- as further evidence of this, Darwin displays an illustration of an orang fetus that regularly
undergoes a stage where the ears are pointed

3) The Age of the Earth

- James Hutton articulated the principle called uniformitarianism that states: geologic
processes taking place now, operated similarly in the past

- relying on this assumption, Hutton and Lyle did empirical studies of the time it takes for
ongoing rock building processes to occur

eg: deposition at beaches and river deltas

eg: accumulation of marine shells (the precursor of limestone)

- Hutton et al. completed a 50 year study to put the rock formations and fossil-bearing strata of
Europe in chronological sequence

- their effort resulted in the geologic time scale

- although Darwin and geologists knew that the Earth was very old, they were unable to precisely
quantify the Earth’s age

- not until radioactive decay was well understood could the Earth’s age be accurately calculated

- it is now known that the Earth is about 4.5 billion years old

Darwin’s Theory of Evolution through Natural Selection

Prior to Darwin’s theory, it was evident that species were not immutable; however, a satisfactory
explanation as to the mechanism of change through time had not been put forth

- in his 1809 Philosophie Zoologique, Lamark proposed that organisms acquire or lose characters
simply by use or disuse

eg: long necks in giraffes became heritable simply because giraffes were stretching their necks
to get to higher leafs

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- that is, behavior somehow created heritable traits

- by this idea a person that spent most of his time flapping his arms like a bird would pass on
wing-like appendages to his offspring

- however, Darwin is now celebrated by biologists because he was the first to understand how
the primary mechanism of evolution works and he also published his theory

- Alfred Wallace formulated and proposed the same theory at the same time as did Darwin;
however, Darwin is more often credited for the theory of evolution through natural
selection because of his more detailed writings

- the primary gap in Darwin’s theory was a lack of understanding of the genetic basis of
evolution

- Darwin states: “The laws governing inheritance are for the most part unknown.”

- this left Darwin’s theory open to criticism well into the 20th century

- although Gregor Mendel published his genetics work in 1866; there is no evidence to suggest
that Darwin ever read Mendel’s works

- this is especially surprising since at the time of Darwin’s death, Mendel’s work was found in
Darwin’s library; however, because Mendel’s work was written in German, perhaps
Darwin didn’t read it

- although Darwin correctly proposed that natural selection was the main driving force in
evolution, he did not discover other forces that drive evolution such as genetic drift

- Darwin did understand the effects of immigration on populations and discussed sexual
selection:

“This form of selection (sexual selection) depends not on a struggle for existence but in relation
to other organic beings or to external conditions but on a struggle between the individual of one
sex, generally the males, for the possession of the other sex. The result is not death to the
unsuccessful competitor but few or no offspring. Sexual selection is, therefore, less rigorous
than natural selection.

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- Darwin’s theory is the logical outcome of the following postulates:

1) individuals within species are variable

2) some of these variations are heritable

3) more offspring are produced than can survive

4) those who survive and reproduce tend to be those that have the most favorable variations (in
the present environment)
ie: survival of the fittest (a term used by Darwin)

- these postulates were laid out in his: On the Origin of Species by Means of Natural Selection or
the Preservation of Favored Races in the Struggle for Life published in 1859

Fitness in this sense means the ability of an individual to survive and reproduce in the current
environment

- an adaptation is a character or trait that increases an individual’s ability to survive and

reproduce in any particular environment

- although an organism may have many features, not all are adaptive

Darwin’s postulates have no hidden assumptions nor do they require uncritical acceptance.

- each postulate is testable via empirical evidence and experiment

- indeed, much of Darwin’s text is concerned with addressing what others considered weaknesses
in his theory

eg: absence of frogs and terrestrial mammals on oceanic islands

- although many oceanic islands are well suited to sustain frogs, they seldom do

- this is because frogs can not survive in sea water like some other organisms can

- terrestrial mammals are likewise rarely found on oceanic islands; however, bats (which can fly)
are common on these islands

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- because terrestrial mammals and frogs typically could survive on these islands, Darwin asks:

“But why, on the theory of creation they (frogs and mammals) should not have been created
there, it would be very difficult to explain.”

- in other words:

1) if special creation had occurred, why were islands not initially stocked with organisms that
were so well adapted to living there, and

2) why are only those organisms on oceanic islands that could have immigrated there on their
own

eg: dispersal of alpine organisms

- the dispersal of alpine organisms on mountain summits was difficult to explain under Darwin’s
Theory

- that is, how could the same plant and animal species be found on mountain summits that may
be separated by hundreds of miles of lowlands where these species could not survive

- Darwin posited that during the glacial period, these alpine species were widespread in the then
cold lowlands, as the glaciers retreated, the species only remained on cold mountain
peaks

- thus, the presence of alpine organisms could be explained without invoking the theory of
special creation

Testing Darwin’s Postulates

The following example demonstrates how each of Darwin’s postulates may be empirically tested:

- Grant & Grant have studied beak size and shape in Galapagos Finches (a group originally
studied by Darwin)

- the 13 species of finches there are similar in size and coloration

- however, they vary by:

1) beak size 2) beak shape

- these beak variations reflect the many different feeding habits among these species

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- this study focuses the medium ground finch, Geospiza fortis

- this is a seed eating species and beak size is correlated with seed size

ie: birds with bigger beaks eat larger seeds

- this situation sets up a good population study because:

1) the species is generally confined to the island Daphne Major

ie: there are few migrations onto or off the island

2) the population is small (about 1,200 individuals) which allows for an exhaustive study
- from the mid 1970's to 1980, virtually all of the population had been captured and marked with
colored aluminum bands

Testing Postulate 1: Are populations variable?

Grant and Grant measured every captured bird for beak length and depth. Additionally, birds
were weighed and their wing and tail length were measured

- when beak depths were plotted, the results formed a bell curve which clearly indicates that
populations are variable

Testing Postulate 2: Is some of the variation heritable?

Variation in characters may be due to environmental factors or genetic factors or both.

- in this example, beak depth could be due exclusively to environmental factors

eg: nutrients received when the birds were chicks could influence the ultimate size of the adult
beak size

- beak depth could also be due exclusively to genetic factors but as in many cases, it is a function
of both

In this study, a colleague of Grant & Grant, Peter Boag estimated a quantity known as

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 heritability

- heritability is the proportion of total variation observed in a population that is due to genetic
factors

- heritability is usually estimated by measuring the similarity between pairs of relatives

- the details of this will be discussed in a subsequent chapter

- however, Boag found a strong similarity between parents and offspring

eg: parents with deep beaks had offspring with deep beaks

eg: parents with shallow beaks had offspring with shallow beaks

- this is evidence that variation in beak depth is under at least some level of genetic control

Testing Postulate 3: Is there an excess of offspring so that only some

individuals survived to mate?
A severe drought on the island in 1977 caused about 84% of the birds to die over a 20 month
period

- most died of starvation and it is clear that only a fraction survived to reproduce the following
year

- although this was an extensive die off, this sort of mortality is apparently not that unusual

- all studies to date suggest that in every natural population, more offspring are produced than
survive to reproduce

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Testing Postulate 4: Is survival and reproduction nonrandom?
By measuring the birds after the 1977 drought, the Grant team determined that birds with deep
beaks survived at higher rates than those with shallow beaks

- therefore the average beak size in the population changed

- during the drought, the large hard fruits of the plant Tribulus cistoides, which are normally
ignored, became an important food source

- typically the birds eat small soft seeds which are available in nondrought years

- only large birds with deep narrow beaks can crack and eat Tribulus fruits

- additionally, large birds defend food sources more successfully

- because large size and deep beaks were positively correlated, the two traits respond to selection
together

In 1983, the island experienced a wet year due to El Ninio. During that year, birds with shallow
beaks were favored due to an abundance of small soft fruits.

- smaller birds were favored over larger birds

- therefore natural selection pressures had changed demonstrating that natural selection is
dynamic

By the criterion that evolution is a response to selection, the change in beak size distribution
between generations demonstrates that evolution had occurred.

- note that the change in beak size distribution within a single generation does not demonstrate
evolution (graph of 1978 survivors)

- it only demonstrates selection has occurred

- if a trait is heritable, then selection will result in an allele frequency change (evolution) within
the population

- therefore, selection and evolution are not the same thing

- why has the beak depth remained large at the end of 1978 but the seed size has returned to
small and soft?

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- can the evolution in beak depth in the birds affect the evolution of the plants that produce the
seeds — yes

- and when this occurs, it is called coevolution

Key Points of Natural Selection

1) Natural selection acts on individuals but its consequences occur in populations

- the effort of finches to crack the large fruits did not result in larger beaks

- instead, the average beak depth and body size in the finch population increased because the
smaller finches died out

- because beak size is at least partly heritable, microevolution occurred because allele
frequencies
in the population had changed

2) Natural selection acts on phenotype but evolution results in changes in allele frequencies

- if body and beak size had not been heritable, selection would have changed phenotype but
would not have cause evolution

eg: if birds had become more muscular during the drought due to territorial defense, then their
phenotypes would have changed but this is not heritable and would not have been passed
on to the next generation

- the idea that phenotypic changes can be passed on the next generation is called Lamarckian
Evolution

- because evolution is the response to selection, it only occurs when traits are heritable

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3) Natural selection is the result of past environmental conditions

- the offspring of the finches that survived the drought will be better adapted to drought
conditions than was the population prior to the drought

- however, they may not be better adapted to the current conditions

eg: a wet year will create new selection pressure on the population

- evolution is always a generation behind environmental changes

- because future environmental conditions are unknown to the forces of natural selection,
organisms can not be adapted for future conditions

4) Natural selection acts on existing traits

- natural selection can only act on traits that already preexist in a population

ie: it can not create a new beak form in response to drought conditions

- however, chance mutations occur continuously in populations and if they are adaptive, they
may
be selected for

- however, natural selection can bring about novel uses for preexisting features

eg: Pandas use a sixth “thumb” to shred bamboo; this is not a true thumb but is a sesmoid wrist
bone

5) Natural selection is directed but not progressive

ie: evolution by natural selection is not a random process nor does it have a predetermined goal

- evolution is directed by the environment and purposeful in the sense of increasing adaptation to
the environment

- the random element in this process is environmental change

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ie: organisms may become more complex or more simple in order to become better adapted to
the environment

eg: the complex mind of humans was not a predetermined goal of evolution

- the increasing complexity of the human mind allowed individuals to defend themselves from
and to hunt animals that were larger and stronger than themselves

6) Natural selection acts on individuals, not populations

- it is a misconception that individuals will do things that are beneficial for the species as a whole

ie: altruistic behavior rarely, if ever, occurs in nonhuman populations

eg: prairie dogs will sound an alarm when predators approach but this occurs only because it
increases the bearer’s fitness relative to competing individuals because the beneficiaries
are close genetic relatives

ie: individuals will provide protection only to those members that share a significant amount of
genetic similarity (close relatives)

- if a gene existed that produced a behavior that reduced the bearer’s fitness and increased the
fitness of others would be strong selected against

The Time Component in Evolution

The small changes in trait distribution and allele frequencies in this example do not explain
differences in traits so great that they may distinguish higher taxonomic levels:

eg: species, genera, families, etc.

- microevolution is the term used to describe small changes that occur within populations

- macroevolution is the term that describes changes necessary to bring about speciation and
higher taxonomic distinction

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- macroevolution requires vast amounts of time which is unimaginable for humans with their
comparatively short life spans

Once rocks have been formed, the radioactive elements in them can be used to date them.

eg: potassium-argon / uranium - lead

- because moon rocks and meteorites are about 4.6 billion years old, the material of the earth is
likely the same age

- however, no rock on earth is quite that old because early earth was molten

- the oldest fossils are cyano-bacteria-like organisms that are about 3.5 billion years old
(stromalite)

- this is an enormous amount of time for evolution to occur

The Evolution of Darwinism

The theory of evolution through natural selection is the organizing theme of all biology.

ie: without evolution, biology is nonsensical

- although Darwinism was introduced to the world in 1859, it took another 70 years before it was
generally accepted, even by biologists

There were 3 serious problems with the theory as originally formulated:

1) the mechanism of variability was unknown

ie: the idea of mutations as the ultimate source of variability was not understood

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2) the mechanism of inheritance was unknown

- it was not until the turn of the 19th century that Mendel’s work was rediscovered and verified

- until then, many biologists (including Darwin) thought that traits were blended like paint
pigments

- Mendel’s work showed that traits were discrete units

- of course some phenotypes appear to undergo blending, but genotypes do not

3) the precise age of the earth was unknown

- at that time, they only credible calculation for the age of the earth was made by Lord Kelvin
who calculated the age at 15-20 million years old

- this age was insufficient to explain the geologic time scale worked out by Hutton & Lyle

- it is evident that Darwin repeatedly struggled with placing biological events within an accurate
time scale

The Modern Synthesis

Not until the mid 20th century did these difficulties become resolved and the fundamentals of
genetics were successfully integrated with Darwin’s four postulates

- this led to reformulation of the theory of evolution and is called “The Modern Synthesis”

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