HL Biology Notes for Evolution

The information in this document covers the IB Biology syllabus for Topic 5.4 and Option D.

Evolution is the process of cumulative change in the heritable characteristics of a population.

Introduction
Evolution involves two interrelated phenomena:
 adaptation: phenotypic variations among members of a species permit some to be more
successful in their environment
 speciation: over the course of time, the number of species (on Earth) multiplies; a single species
can evolve into two or more new species

Charles Darwin, in his book On the Origin of Species (1859), stated that all living things on Earth today
are descendants, with modifications, of earlier species. Darwin believed that all species are related,
meaning that any two species on Earth had a common ancestor (though you may have to go very far
back in time to find a common ancestor between elephants and algae). He proposed a mechanism,
which he called Natural Selection, to explain how new species evolved.

Darwin's Influences
 Erasmus Darwin (Charles Darwin's grandfather) proposed a theory of evolution by “the
inheritance of acquired characteristics”.
 John Baptiste de Lamarck was the first person to publish a reasoned theory of evolution
(Philosophe Zoologique); proposed the idea of use and disuse and inheritance of acquired
characteristics.
 Rev. Thomas Malthus wrote An Essay on the Principles of Population, attempting to justify the
squalid conditions of the poor by stating that poverty and starvation were merely a consequence
of overpopulation.
 Charles Lyell's work Principles of Geology proposed that the geological processes that we
observe today have always been occurring and the Earth is therefore very old.
 Charles Darwin spent five years aboard the HMS Beagle, travelling the world as ship's
naturalist. He observed many phenomena including variation among finches and tortoises on
the Galapagos Islands:

http://faculty.southwest.tn.edu/rburkett/GB%20Pro2.jpg

 Alfred Russell Wallace had also been influenced by Lamarck and Erasmus Darwin, and was a
catalyst in Charles Darwin's actually publishing The Origin of Species.

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HL Biology Notes for Evolution

Why Natural Selection?

Darwin had observed that in domesticated animals (such as dogs), breeders selectively bred animals with
favoured characteristics – artificial selection. He proposed that a natural phenomenon, which he called
“natural selection”, had resulted in the development of new species over time.
The main points of Darwin's theory of evolution by natural selection are as follows:
 living things produce more offspring than the finite resources available to them can support
 thus living things face a constant struggle for existence
 the individuals in a population vary in their phenotypes (appearance) – this variation is a result of
sexual reproduction
 some of this variation is heritable – it is a reflection of variation in genotypes (DNA)
 those variants best adapted to the conditions of their life are most likely to survive and reproduce
themselves – survival of the fittest
 to the extent that their adaptations are heritable, they will be passed on to their offspring

The forces of natural selection act on phenotypes, but only if there is a change in the genotypes of a
population has evolution occurred.
How is “Fitness” Measured?
Fitness is a measure of reproductive success. Those individuals who leave the largest number of mature
offspring are the fittest. This can be achieved by:
 Survival/mortality selection – any trait that promotes survival, at least until the reproductive years
are over, increases fitness.
 Sexual selection – one gender, usually the females, chooses among available mates. Any
inherited trait that improves the mating success of certain individuals will become more
pronounced in successive generations.
 Fecundity selection – the production of a large number of mature offspring is a measure of fitness.
(Only mature offspring can pass on traits to the next generation.)
If a species cannot adapt to the changing environment, then the species will die out. As the dinosaurs
(cold-blooded) did not find a way to adapt to a cooler climate, they did not survive, and the warm-
blooded mammals became the dominant animal species on Earth.
Natural selection leads to the increased reproduction of individuals with favourable heritable variations.
Variation is non-directional and/or random. The selection process is dictated by the environment and
leads to differential survival. This results in survival of those individuals best adapted to their particular
environment – they will be able to get the most food, find the best shelter, find a mate, and reproduce.
They will also be best able to avoid predators and care for their offspring.

Examples of Natural Selection in Response to Environmental Change

 Bacteria cause many illnesses in humans. Antibiotics are drugs which block specific steps in
bacterial metabolic pathways, resulting in death of the bacteria. Since World War II, many
bacteria have become resistant to antibiotics, and the former “wonder drugs” are now ineffective
against certain strains of bacteria. For example, Staphylococcus aureus is a common bacterium
found living on skin. It is usually harmless, but under some circumstances can enter the
bloodstream, infect tissues in the kidneys or bones, and could become fatal. There are some
strains of S. aureus that are resistant to all known antibiotics. These bacteria are of grave concern
to hospitals, where patients have died of a simple Staph infection, once treatable by antibiotics.

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• Pesticides are substances used for destroying insects or other organisms harmful to cultivated
plants or to animals. Pest populations have evolved resistance to these chemicals, and, as such,
they are losing their effectiveness. This can be seen as an example of artificial selection, as
humans have caused it. For example, millions of rats aren't even getting sick from pesticide
doses that once killed them. In one county in England, these "super rats" have built up such
resistance to certain toxins that they can consume five times as much poison as rats in other
counties before dying.

Types of Evolution:

• Co-Evolution occurs when two or more organisms evolve together. Three types of co-evolution
are predator-prey relationships, symbiotic relationships, and parasitic relationships.
• Microevolution is a change in a population's alleles over a period of time.
• Macroevolution is the changes that occur over a long period of time, resulting in new species that
replace old species.

Convergent Evolution occurs when species with different ancestors evolve to look similar as a result of
adaptation to similar environments. For example, dolphins (mammals) and sharks (fish) have similar
body types because they live in similar environments.

http://www.hickerphoto.com/data/media/45/dolphin-photos_4786.jpg http://en.wikivisual.com/images/2/24/Oceanic_Whitetip_Shark.png
Divergent Evolution occurs when two or more similar species become more and more dissimilar as a
result of environmental adaptations. For example, red foxes live in the forest, and kit foxes live in the
desert, and have evolved differently as a result. Adaptive radiation is an example of divergent evolution.

Red Fox
Kit Fox (images from www.canids.org)

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 During his voyage on the Beagle, Darwin observed that finches on the Galapagos Islands show
great variations in the size and shapes of their beaks. These birds had been isolated from the rest
of the world for a long period of time. Their beak size and shape is directly related to the type of
food they eat:

http://www.greenclub.bc.ca/Green_Club_Activity/GVRD_Parks_Events/Finches.jpg

The Origin of Life on Earth

Before there were living things on our planet, the Earth was a very different place from what we now
recognize as “home”.
 The atmosphere most likely contained H2, CO2, NH3, CH4, and water vapour. It is considered to
have been a “reducing atmosphere” because metals contained in old rocks are in their reduced
form (e.g. iron (II) vs. iron (III), which is the oxidized form).
 There were frequent thunderstorms (lightning), and no ozone, which meant high levels of UV
radiation.
There are a few theories that attempt to account for the origin of life on Earth:

1. Panspermia is the theory that life originated somewhere other than Earth (i.e. from space).
• On September 28, 1969, a meteorite fell near Murchison, Australia. It was found to contain a
variety of organic molecules:
 purines and pyrimidines
 polyols such as glycerol and sugars
 amino acids (90 different structures, including 19 that are found on Earth)
• The ALH84001 meteorite arrived here from Mars. It contained a variety of organic molecules
and evidence of microorganisms (bacteria).

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2. Special Creation mostly refers to the origin of humans. Many religions say that God created
humans. As a result many people feel that the concept of evolution goes against their religion. It
can also refer to the belief that a Supreme Being created all of Earth, including all living things.

3. Spontaneous Generation of life would have required four processes to occur.

• non-living synthesis of simple organic molecules such as sugars and amino acids
• polymerization, in which these molecules are joined to form biological macromolecules
• origin of self-replicating molecules that make inheritance possible (e.g. DNA & RNA)
• packaging of these molecules into membranes with a different
internal chemistry than the external environment

Stanley Miller and Harold Urey investigated Spontaneous

Generation in 1953. They set up an apparatus similar to the one at
right:
 Miller hypothesized that the mixture of gases he used (no
oxygen!!) resembled the atmosphere of the early Earth. The
spark simulated lightning.
 By continuously boiling and condensing the water, the mixture
was kept circulating.
 At the end of a week, several amino acids and other organic
molecules were present.
 When this experiment was repeated with the addition of CO2, a
greater variety of organic molecules were formed.

In 1961, Juan Oro was able to make amino acids from an aqueous solution http://users.rcn.com/jkimball.ma.ultranet/
BiologyPages/M/Miller.gif
of ammonia and hydrogen cyanide. He also made large quantities of
adenine, an important component of RNA, DNA and ATP.

One problem: all metabolic pathways depend on enzymes, which are made of protein.
 Proteins cannot be made without nucleic acids.
 Nucleic acids cannot be made without proteins (i.e. enzymes).
The discovery that certain RNA molecules have enzymatic activity provides a possible solution:
 Ribozymes incorporate both
features required of life:
information storage and
catalytic ability. The
ribozyme serves as both the
template on which short
lengths of RNA -
oligonucleotides – are
assembled following the rules
of base pairing and the
catalyst for covalently linking
these oligonucleotides.
http://users.rcn.com/jkimball.ma.ultranet/BiologyPages/S/Szostak.gif

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 Where did the ribozymes come from? The catalytic action of clay is a suggested mechanism:
■ Some kinds of clay can grow by attracting molecules to themselves. They will then repeat
a lattice-like structure over and over again.
■ Amino acids may have stuck to the clay lattice and have been incorporated into it. They
may have been attached to each other as well.
■ Some clay particles may have become a template for a protein.
■ If the protein product was a weak enzyme it may have sped up the process of “protein
synthesis” with clay as a template.
■ Then the clay template for this particular protein would make more protein than another
template whose product was not an enzyme.
■ Then nucleotides could have become attracted by the clay template, or the template with
the attached proteins, and could have polymerized (into RNA) and come to act as a co-
enzyme.
■ The more successful template is the one where the enzyme and
co-enzyme work together to produce more for themselves.
■ Eventually the co-enzyme could become the template for protein
synthesis – and a ribozyme is born!

Ribosomes, the organelles in cells responsible for protein synthesis, are an

example of a ribozyme. It is composed of rRNA (ribosomal RNA) and catalyzes
the polymerization of amino acids into polypeptide chains.

Where Did Organic Molecules Originate?

• In space: spectral line analysis of clouds of cosmic dust particles has revealed presence of
glycine, a simple amino acid. Combining this evidence with that of meteorites suggests that the
first organic molecules could have come from an extraterrestrial source.
• In alternating wet/dry conditions: the rise and fall of tides results in a
continuing cycle of wet/dry conditions in the intertidal zone. This also
occurs in the flood plains of rivers. Drying of clay particles in these
zones could have contributed to polymerization of amino acids and
nucleotides. Ancient organisms called stromatolites (right) found living
in intertidal zones support this theory.
• Near volcanoes: when volcanoes
erupt, they release water vapour, other gases and minerals that
could be used to form organic matter. The combination of raw
materials and heat could have provided the conditions
necessary to form simple organic compounds.
• Hydrothermal Vents: deep in the oceans,
hydrothermal vents are places where hot
water spreads out from beneath the ocean floor. As the hot water rises, it
picks up minerals from the Earth's crust. There are entire communities of
organisms living near these "black smokers" (right), which not only disproves
the idea that there can be nothing living so deep in the oceans, but that this
could well be where organic molecules first formed.

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The Origin of Cells

We know that cellular membranes are phospholipid bi-layers with proteins. Their main function is to
separate the inside of the cell from the outside environment, and to control the passage of materials into
and out of the cell.
 In 1957, Sidney Fox heated amino acids in the absence of water. The amino acids polymerized
into protein chains.
 When water was added and the mixture cooled, small, stable microspheres (a.k.a.
coacervates) were formed. These are both examples of protobionts.
 The microspheres accumulated substances inside themselves and attracted lipids to the
outside, forming a lipid-protein layer.
 The combination of these microspheres with ribozymes would have formed very primitive
cells. It is believed that this occurred 3.8 b.y.a..

Alexander Oparin suggested that different types of coacervates might have formed
in Earth's primordial sea and subsequently been subject to natural selection,
leading to the origin of Life. Eventually, prokaryotes evolved.

commons.wikimedia.org
What about Oxygen?

In the early Earth, there was very little

oxygen in the atmosphere. Most of it was
locked up in compounds. Water vapour in
the atmosphere can be split by UV radiation
– this would account for the limited
quantities of oxygen in the atmosphere at
the time. Around 2.5 b.y.a., after
prokaryotes had existed for nearly two
billion years, a form of chlorophyll appeared
in
bacteria that allowed photosynthesis –
which we know releases O2.

The rising oxygen levels allowed for certain

changes to the Earth's chemistry:

• breakdown of compounds into

oxidised mineral sediments and
carbon dioxide
• formation of the ozone layer, which filtered out UV radiation and stopped production of the
more noxious compounds in the atmosphere

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The Origin of Eukaryotes = Endosymbiotic Theory

In 1966, Lynn Margulis proposed that mitochondria were once independent organisms.
These pre-mitochondria were engulfed by a bacterium (by phagocytosis) and formed a
symbiotic relationship with the host cell, receiving food from the host, and supplying the
host with ATP. Chloroplasts formed when a cell that had mitochondria engulfed an
autotrophic cell.

Evidence for the Endosymbiotic Theory:

 chloroplasts and mitochondria have a double membrane, which would have been formed by the
process of phagocytosis
 chloroplasts and mitochondria have their own DNA and ribosomes
 ribosomes and DNA in chloroplasts and mitochondria are very similar in structure to bacterial
ribosomes and DNA

Evidence for Evolution

• Comparative Anatomy looks at similarities
and differences among the physical structures
of organisms. One key piece of evidence for
evolution is the pentadactyl limb of
vertebrates:

http://apbio12007.blogspot.com/2007/09/evidence-of-evolution-by-
natural.html

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Homologous Structures Analogous Structures

• are similar in fundamental structure • resemble each other in function
• are similar in position and development • differ in their fundamental structure
• are similar because of common ancestry • illustrate only superficial resemblances
• e.g. pentadactyl limb of vertebrates • e.g. wings of insects, bats and birds

 Comparative Embryology looks at the embryonic development of different species. In the early
stages of life, there are many similarities among vertebrates:

http://apbio12007.blogspot.com/2007/09/evidence-of-evolution-by-natural.html

• Selective Breeding of Domestic Animals

o Artificial selection is the process by which all plants and animals used by humans have
been derived from wild organisms. It involves identifying the largest, best or most useful
of the progeny for the intended purpose and using them as the parents of the next
generation.
o By “selecting” those traits most desired, the genetic makeup of a population changes
rapidly.
 Pomeranians, Pugs, Keeshonds and Great Danes are all examples of Canis
familiaris, show obvious differences that make us question their classification.

Images from http://www.justdogbreeds.com

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• Fossils are the preserved remains or traces of prehistoric life.

 Petrified wood or bone (turned to stone when internal cavities and pores are filled
with mineral matter)
 Impressions such as leaf prints, footprints, and the like
 Coprolites (petrified feces)
 Insects in amber (hardened tree sap)
 Frozen mammoths in the Arctic
 Mummified corpses in peat bogs
o The fossil record is incomplete, however, there is evidence to show how some species
have evolved:

Image from http://www.sanparks.org/parks/kruger/elephants/about/evolution.php

o Relative age of fossils can be
determined by looking at the
sedimentary layers each is found in –
those closer to the surface are
assumed to be more recent than
those deeper in the Earth.
o Rocks and fossils can also be “dated”
using radioisotopes of carbon (C-14)
and potassium (K-40):
 half life is the fixed interval in
which half of the amount of a
radioactive substance will
decay
http://www.splung.com/nuclear/images/radioactivity/half-life2.png

 Carbon dating has become an excellent method of determining age of organic

material. In living tissues, some of the carbon present will be the radioactive
isotope, C-14. Most of it will be the “normal” form, C-12. Its nucleus contains 6
neutrons & 6 protons, and has 6 electrons in orbitals. C-14 has two extra
neutrons, which makes it relatively unstable – it will spontaneously change into
N-14, emitting radiation as it does so. As long as a cell is alive, it will be
constantly replenishing its supply of C-14 – after death, it will diminish, which
allows us to use the proportion of C-12 to C-14 to estimate the age of a fossil.
• The half-life of C-14 is 5730 years – this means that 5730 years after the
death of an organism, the amount of radioactive C-14 present will be half
of what it was when the organism was alive.
• Carbon dating is useful for estimating the age of materials between 20 000
and 50 000 years old.
• For rocks, which are often much older than 50 000 years, we can use K-

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40, which has a half-life of 1300 million years.

o Paleontological evidence for evolution is based on fossils found. There are fossils of horse
ancestors dating back to 53 m.y.a., and many intermediate species in between.

• Biochemical Evidence uses common molecules found in organisms.

o DNA, as we know, is found in all organisms. The genetic code is universal, and all
organisms use the same 20 amino acids to make proteins.
o Comparing amino acid sequences in similar proteins from different species (e.g.
hemoglobin, cytochrome c) can give an indication of how long ago the species diverged
from a common ancestor.
o Biochemical evidence is generally used together with other kinds of data such as
paleontological data.

Phylogeny is the line of evolutionary descent.

 Biochemical evidence can be used to indicate phylogeny – this requires sequencing specific
proteins from different species:
 Analysis of hemoglobin in primates has suggested that humans are more closely related to
chimpanzees than other apes – this tells us that the common ancestor between humans and
chimps was more recent than the common ancestor we share with gorillas.
 Cytochrome c, a protein in the electron transport chain, is often used for less-related species
■ cytochrome c in chimpanzees and humans is identical, and only differs in one amino
acid from that of a rhesus monkey
■ comparing cytochrome c between humans and bacteria or yeast has also shown that
there are similar sequences between the proteins

Cladistics is the analysis of how species can be grouped into clades. It is used to determine the relative
timing of species divergences.
i. Cladogram: a diagram depicting patterns of shared characteristics among species.
ii. Clade: a group of species that includes an ancestral species and all its descendants.

Monophyletic group = a clade Paraphyletic group ≠ a clade Polyphyletic group ≠ a clade

Images from https://eapbiofield.wikispaces.com/Chapter+26+Phylogeny+AJ

Assumptions in Cladistics:
• The group of organisms is related by descent from a common ancestor.
• Descent follows a bifurcating pattern where the ancestral form splits into two sister taxa and the
ancestor goes extinct.
• Change of characteristics occurs in lineages over time.

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Climbing the Human Evolutionary Tree

Humans and other primates share a common

ancestor:

The first primates were small arboreal (tree-

dwelling) mammals. Primates originated 65
m.y.a., and are defined by characteristics which
make them adapted for living in trees.

© Gary Larson

http://users.rcn.com/jkimball.ma.ultranet/BiologyPages/P/PrimatePhylogeny-rev3.tif
Modern Primates are divided into two sub-orders:
 Prosimians are the most primitive primate species, and are believed to have diverged from the
other primates more than thirty million years ago.

lemurs lorises pottos tarsiers

 Anthropoids include monkeys, apes and humans.

 Monkeys include the New World monkeys (found in the Americas) and Old World monkeys
(found in Africa & Asia). These groups evolved along separate pathways.

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New World Monkeys Old World Monkeys

Above: Douc languor

Above: Capuchin
Right: Mandrill
Right: Golden Lion Tamarin

 Apes are confined to tropical regions of the world. They are larger than monkeys and have
longer arms and no tail. They also have larger brains than monkeys. There are four genera of
apes:
■ Hylobates = gibbons ■ Gorilla = gorillas
■ Pongo = orang-utans ■ Pan= chimpanzees

gibbons orang-utans gorillas chimpanzees

 Humans are most closely "related" to chimpanzees, and have no other living "relatives". Of
the two species of chimpanzees, humans are most closely related to the Bonobos.
Human History
The study of human origins is called paleoanthropology. There have been several misconceptions about
the origin of humans, including descent from chimpanzees or other modern apes. Current thinking
suggests that the human evolutionary tree includes many dead end branches, and that there were several
species of humans in existence at the same time.
Ardipithecus ramidus is considered to be the earliest member of the Hominoidea family; this is because it
is the most ape-like hominid known. Timothy White discovered the first fossils of this species in Ethiopia
in 1992. It existed 4.4 million years ago (m.y.a.), and was about the size of a modern chimpanzee.
The Australopithecines are considered to be the first humans, and are still quite ape-like in their
characteristics:
 Raymond Dart discovered Australopithecus africanus (Taung child). It walked upright, had
human-like teeth, and had a brain about 1/3 the size of modern humans. They appeared about
4.4 m.y.a., and existed for about three million years.
 Australopithecus afarensis was a close relative of A. africanus. The best-known example of this
species is "Lucy", discovered in Ethiopia. They walked upright, and were similar in size to a
chimpanzee. They lived from 3-2 m.y.a..

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 Australopithecus robustus lived from 2.4-1.4 m.y.a., and were larger and more heavily built than
older species.
Members of the genus Homo more human-like than the Australopithecines:
 Mary and Louis Leakey discovered Homo habilis ("handy man") in Tanzania. Fossils of this
species show evidence of a larger brain, and were found in the same location as simple stone
tools. H. habilis lived about 2.5 m.y.a., coexisting with Australopithecines.
 Homo erectus ("upright-walking man") are believed to have originated 1 m.y.a., in Africa. They
were the first hominid species to leave Africa, spreading into Europe and Asia. H. erectus had a
larger brain than H. habilis, and made more complex tools.
 Homo ergaster ("working man") is a name given to early forms of Homo erectus (pre-migration),
and are characterized by a rounded cranium, a brow ridge, and smaller teeth than
Australopithecines.
 Homo neanderthalensis descended from H. erectus, spreading into Europe, the Middle East and
Asia. They were adapted to a colder climate, with a short, stocky shape, and a large nasal
opening, which would have given them a keen sense of smell. Many Neanderthal sites have
evidence of purposeful burials, and other signs of cultural development.
 Homo sapiens is the modern human, the earliest of which is Cro Magnon man. Cro Magnon had
a close resemblance to living humans. They are suspected to be the first to use language.
There is still disagreement among the
scientific community about the
classification of the more recent Homo
species, including a suggestion that H.
sapiens and H. neanderthalensis arose
from H. heidelbergensis, once called
Archaic Homo sapiens.

Classification of Humans
 Kingdom Animalia
 Phylum Chordata
 Subphylum Vertebrata
 Class Mammalia
 Subclass Eutheria
 Order Primates
 Suborder Anthropoids
 Family Hominidae
 Genus Homo
 Species sapiens
 Subspecies sapiens

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What makes Humans primates?

 Opposable Thumbs
 improves ability to grasp objects (e.g. tree branches)
 allows manipulation of objects (e.g. tools)
 Acute Vision
 eyes at front of head gives a smaller field of vision but allows for distance judgement
■ vital for swinging from tree to tree
■ colour vision enhances chances of locating food
 Large Brain
 improves ability to manipulate objects
 related to long-term care of young
 Extended Care of Offspring
 babies are born helpless
■ modification of pelvis and birth canal requires that the baby's head is smaller
 gives opportunity to learn many skills from parents
 Teeth
 having different types of teeth allows primates to have a varied diet (omnivores)

Humans are a bipedal and neotenous species of African ape that spread to colonize new areas.
 Bipedal = walking on two legs
 Neotenous = having extended care of young

Evidence:
 Anatomy
 Homo erectus is believed to be our direct ancestor.
Fossil skeletons indicate that their body type was very
similar to ours. (The skull, however, is still very ape-
like.)
■ Homo erectus originated in North Africa two
million years ago, spreading through Europe and
Asia.
 Tool Use
 H. erectus used purpose-made tools. Early hominids such as “Lucy” used tools of
opportunity.
 Biochemistry
 Mitochondrial DNA (mtDNA) is passed along maternal lines, since the only paternal
contribution to the zygote is DNA. Because mtDNA changes more rapidly than nuclear
DNA, it is useful for evolutionary studies.
 Studying the mitochondria of different groups of people (European, African, etc.) has shown
that there is more variation in mtDNA among Africans than all other humans.
■ Homo sapiens existed in Africa 200 000 years ago.
■ They moved to other parts of the world about 100 000 years ago.
■ Therefore, H. sapiens remaining in Africa had more time to develop different kinds of
mtDNA.

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Ecology of Early Humans

Fossil evidence has indicated that as early as 3.7 m.y.a., humans were walking upright.
 fossil footprints in volcanic ash
 skeletal evidence (e.g. “Lucy”)

Bipedalism meant that less surface area was exposed to the sun's most intense rays, an advantage on the
plains of Africa.

As the climate cooled, savannas replaced forests – this meant fewer trees to climb. Hominids climbed
trees less, and bipedalism freed up the hands for tool use. Increase in brain size lead to more advanced
tool making, and made early hominids better able to deal with the changing environment.

Homo species replaced Australopithecus since they were less dependent on trees.

A drier climate meant it was more difficult to find edible vegetation.

 Individuals who supplemented their diet with meat were better able to survive, and the protein
aided brain growth.
 Less dependence on familiar plants as a source of food allowed for migration to new areas.

Neanderthals had short, thick bodies that retained heat better than the taller, slimmer form of Homo
erectus.
 Neanderthals were better adapted to the colder climate of Europe.
 Their powerful muscles allowed them to hunt larger animals.

It is believed that modern humans supplemented their diets with fish, leading to increased brain growth.
This is likely why Homo sapiens eventually replaced the Neanderthals.

Cultural Evolution
Genetic evolution involves the change of genetic material, which is subsequently passed on to future
generations. Genetic change is usually random, and the environment dictates whether or not the change
is favourable. Skills acquired by an organism are not passed on to its offspring genetically – the child of
a great hunter would have to be taught how to hunt.

Cultural evolution is the accumulation of useful skills and knowledge and the discarding of harmful
practices that have been handed down from generation to generation for thousands of years. It has
occurred in humans because we have language. Language can be used to pass accumulated knowledge
to offspring, so that it does not need to be re-learned. The evolution of human speech occurred at the
same time as the emergence of the reflective mind, or consciousness, in the Homo lineage.
Brain Size
There is significant evidence that increase in brain size is related to diet. Since the brain is an
“expensive” organ, requiring a lot of energy to operate, and Australopithecus was primarily a vegetarian,
requiring large volumes of food to fulfil energy requirements, its brain was proportionally smaller than
early species of Homo, which ate more meat.
Tool Use
Use of tools by early humans coincides with the increase in brain size. The oldest fossil tools are 2.6
m.y.a., however, there are no fossils of Homo that are this old, which may indicate that either

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Australopithecus used tools, or another, earlier form of Homo existed that we have not found fossils of. It
may also be that an intermediate species, no longer Australopithecus, but not yet Homo existed.
Religion
It is believed that Neanderthals had a religion – they buried their dead with flowers and tools, which
suggests a belief in an afterlife. Earlier species of Homo did not even bury their dead.
Art
Neanderthals did not leave behind much evidence of art, however decorative beads have been found
with Neanderthal fossils. Cro Magnons (early H. sapiens) painted drawings of animals on the walls of
caves, and had elaborate art that was possibly related to a form of religion.
Agriculture
Cultural evolution led to the domestication of animals and agriculture approximately ten thousand years
ago. This allowed humans to form settlements, rather than their previous hunter-gatherer societies in
which they followed their prey, rather than settling in one place.

Species & Speciation

Darwin and Wallace published their evolutionary theories before knowledge of the mechanism of
inheritance – genetics. Gregor Mendel's work was not published until after the Origin of Species. Neo-
Darwinism takes into account both the force of Natural Selection and the mechanism of genetic
inheritance and variation.

Population genetics is the study of the genotypes and their changes in entire populations.
 A gene pool consists of all the genes and their different alleles, present in an interbreeding
population.
 Allele frequency is the commonness of the occurrence of any particular allele in a population.

Evolution involves a change in allele frequency in a population’s gene pool over a number of
generations.

Barriers between gene pools prevent interbreeding between species, and can result in speciation.

Geographic Isolation occurs when what was once a continuous population is divided into two or more
smaller populations.
1. This can occur when rivers change course, mountains rise, continents drift, or organisms migrate.
2. Humans can cause this by building highways, pipelines, or even fences.

http://evolution.berkeley.edu/evolibrary/article/evo_43

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The physical barrier keeps them from mating with one another (reproductive isolation), and differences
from one environment to the other, along with variations in allele frequency between the groups, can
lead to speciation.

Reproductive Isolation

After geographic isolation or another mechanism separates a population into two groups that do not
interbreed, there need to be internal mechanisms to prevent gene flow if the species do mix again -
otherwise, speciation is not complete. Reproductive isolating mechanisms are either pre-zygotic,
preventing copulation, or post-zygotic, resulting in no further reproduction of any offspring formed.
1. Behavioural isolation includes differences in courtship rituals and mating displays. If the
behaviour is too different, the intended mate won't be interested.
2. Temporal isolation occurs when mating season differs among related species. For example, two
species of toad, both the same genus (Bufo), have mated in a laboratory setting to produce
offspring, but in the wild, one mates only in early summer, the other only at the summer's end.
3. Structural isolation occurs when there are physical barriers to mating, such as incompatibility
between sexual organs, as illustrated by the damselfly penises below:

http://evolution.berkeley.edu/evolibrary/article/evo_44

Inter-Species Hybrids

In the event that individuals from different species do mate, there are post-zygotic mechanisms that
prevent success of the offspring.
• Hybrid inviability occurs when an offspring is produced, but does not survive gestation. A hybrid
between Ovis aries (sheep, 2n = 54) and Capra hircus (goat, 2n = 60) is usually stillborn.
• Hybrid infertility occurs when a viable offspring is produced, but it is sterile. Mules are the
infertile offspring of the cross Equus asinus (male donkey, 2n = 62) x Equus caballus (female
horse, 2n = 64). Since mules have 63 chromosomes, an odd number, they cannot form gametes.

Defining 'Species'

The dictionary definition of 'species' is: a group of living organisms

consisting of similar individuals capable of exchanging genes or
interbreeding. The species is the principal natural taxonomic unit, ranking
below a genus and denoted by a Latin binomial, e.g., Homo sapiens.

When dealing in evolutionary terms, we also have a different definition of

species: a part of a potentially interbreeding population with similar
DNA. They have a common gene pool.

Ecologically, we can define a species as a group of organisms adapted to

a particular set of resources in the environment - what we call niche.

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This brings into question some of the aspects normally used to define members of a species:
 the organisms should look alike
 the organisms should interbreed
 the organisms should produce fertile offspring

If these criteria are met, one could consider organisms to be of the same species. However, there are
situations that exist where strictly using these criteria can cause problems:

• Organisms that interbreed do not always look alike:

o Lions and tigers can interbreed, producing a liger or a
tigon. Some ligers and tigons are fertile. Does this mean
that lions and tigers are the same species?When two
organisms of different species interbreed, they produce an
interspecific hybrid. These are usually sterile (as are mules,
normally). If the offspring are fertile, then by the above
criteria, the two parent organisms should be considered the
same species.
• We cannot use the second and third criteria when discussing extinct species.
o Without genetic analysis, it was difficult to determine whether Neanderthals were sub-
species of Homo sapiens, or a separate species.

• Some species reproduce solely by asexual means. This also negates

use of the second and third criteria.
o Some bacteria species can reproduce sexually by conjugation,
exchanging plasmids with other bacteria – they do not have
to be the same species for this to occur.
o
• Ring species have a geographic distribution that
forms a ring and overlaps at the ends.
o The many subspecies of Ensatina
salamanders in California exhibit subtle
morphological and genetic differences all
along their range. All of them interbreed
with their immediate neighbours with
one exception: where the extreme ends
of the range overlap in Southern
California, E. klauberi and E. eschscholtzii
do not interbreed. So where do we mark
the point of speciation?

http://www.ucmp.berkeley.edu/blog/wp-content/uploads/2010/03/Ensatina-slide-400_web.jpg
The existence of these problems does not require us to reject the definition of species, or the criteria used
to define them – they work for most cases – but it is necessary to be aware that there may be situations in
which the species definition is difficult to apply.

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Polymorphisms
A polymorphism is the occurrence of different forms among the members of a population or colony, or in
the life cycle of an individual organism.

When different forms coexist within a given population in equilibrium, we call this a balanced
polymorphism. The forms are present in a constant proportion from generation to generation.
• In humans, sickle cell disease is an example of a balanced polymorphism. Individuals who are
heterozygous for the sickle allele have an advantage in areas where malaria is common.

Sickle Cell Distribution Normal & Sickled RBC's Malaria Distribution

A transient polymorphism occurs when one form is gradually replacing another. It is a temporary
situation that occurs as a by-product of directional selection.
 The peppered moth, Biston betularia, occurs in two forms (morphs). One is grey with dark
splotches, and one is completely dark (melanic). The grey (mottled) form is well-camouflaged
against the lichen-covered bark of trees in unpolluted regions, which protects them from
predators. In this environment, the melanic form is at a disadvantage.
 During the Industrial Revolution in England, the air quality declined, killing off lichens and
resulting in a marked increase in the frequency of dark moths – they stood a better chance
against the dark tree bark than the light-coloured lichens.

Melanic form camouflaged on dark background Mottled form camouflaged on light background

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Allopatric vs. Sympatric Speciation

Allopatric speciation is the evolution of one species into another in different geographic areas.
• A 'break-away' population can be isolated during
migration. For example, a storm could blow a group of
migrating birds off their route, or a small group could be
left behind when the main population migrates.
• Allopatric speciation occurs in island chains - for
example, the Galapagos Islands.

Experimental Evidence for Allopatric Speciation:

In 1989, Diane Dodd conducted an experiment that

demonstrated allopatric speciation in the fruit fly (Drosophila
pseudoobscura):
• A single population of flies was divided into two, with
one of the populations fed with starch-based food and
the other with maltose-based food.
• After the populations had diverged over many
generations, the groups were again mixed; it was
observed that the flies would mate only with others from
their adapted population.
http://evolution.berkeley.edu/evolibrary/article/evo_45

Sympatric speciation is the evolution of one species into another in the same geographic area.
Reasons for reproductive isolation could be ecological (exploiting a new niche) or behavioural
(differences in mating calls or migratory behaviour).

An example of Sympatric Speciation:

Apple maggot flies originally laid their eggs only on hawthorns—but today, these flies lay eggs on
hawthorns (native to North America) and domestic apples (introduced to North America).
• Females generally lay their eggs on the type of fruit they grew up in, and males look for mates on
the type of fruit they grew up in.
• Hawthorn flies generally end up mating
with other hawthorn flies and apple flies
generally end up mating with other apple
flies.
• This means that gene flow between parts of
the population that mate on different types
of fruit is reduced. This host shift from
hawthorns to apples may be the first step
toward sympatric speciation—in fewer than
200 years, some genetic differences
between these two groups of flies have
evolved.

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Polyploidy
Speciation by polyploidy is an example of sympatric speciation. It occurs almost exclusively in plants.

A ploidy change generally means multiplying the number of chromosomes the species has by some
number. A species that normally has 18 chromosomes might produce a lineage that has 36 or 54
chromosomes. Ploidy changes often produce a species that is reproductively isolated and distinct from
the “parent” species. For example, speciation in these anemones involved a ploidy change.

A cross between these two flowers

(2n x 4n) would produce sterile
triploid (3n) offspring, unable to
undergo meiosis because of the odd
number of chromosomes.

Hardy-Weinberg Principle
When the gene pool of a population remains more of less unchanged, then we know that the population
is not evolving. We can detect changes in the frequency of alleles (and thus changes in the gene pool)
using the Hardy-Weinberg formula.

The general formula representing allele frequencies (two alleles) is:

0 < p < 1 and 0 < q < 1 p+q=1 p = frequency of dominant allele

q = frequency of recessive allele

The Hardy-Weinberg formula is used to describe stable gene pools:

p2 + 2pq + q2 = 1
p2 = frequency of
q2 = frequency of
homozygous
homozygous
dominant phenotype
recessive phenotype

2pq = frequency of
heterozygous
phenotype

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Conditions for Using the Hardy-Weinberg formula

There are certain assumptions when using the Hardy-Weinberg formula:
• The population size is very large.
• Mating is random.
o No sexual selection.
o Any individual has an equal chance of mating with another individual.
• No mutations are occurring.
• There is no immigration or emigration (i.e. a closed population).

Example:
In Canada, the genetic disorder cystic fibrosis occurs in 4/10 000 persons. Use the Hardy-Weinberg
Principle to calculate the frequency of the cystic fibrosis allele.

Let "p" represent the normal allele and let "q" represent the cystic fibrous allele.
p+q=1

Individuals with cystic fibrous are homozygous recessive, or q2 and q2 = 4/10 000 persons or 0.0004.

q2 = 0.0004, so q = 0.02

p + q = 1, so p = 1 – q = 1 – 0.02 = 0.98

2pq + q2 gives the number of individuals in the population that carry the allele:

2(0.98)(0.02) + (0.0004) = 0.0396

The cystic fibrosis allele has a frequency of 2%; approximately 4% of Canadians carry the allele.

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