PLB 201- EVOLUTION AND SPECIATION- Dr S.B.

Adeyemi
Evolution has been described as series of changes across successive generations in the heritable
characteristics of biological populations. Processes in evolution give rise to diversity at every level
of biological organisation, which includes species, individual organisms and at the molecular level,
such as proteins and DNA. Approximately 3.7 billion years ago, life on Earth originated and then
evolved from a universal common ancestor. The divergence and repeated speciation of life can be
traced through shared sets of biochemical and morphological traits, or by shared DNA sequences.
Existing patterns of biodiversity have been shaped both by speciation and by extinction. The theory
of evolution by natural selection was formulated by Charles Darwin, who was the first to recognise
natural selection as an important cause of evolution.
Evolution Pre-Darwin
Contrary to many assumptions, evolutionary theory did not begin in 1859 with Charles Darwin
and The Origin of Species. Rather, evolution-like ideas had existed since the times of the Greeks,
and had been in and out of favour in the periods between ancient Greece and Victorian England.
Indeed, by Darwin's time the idea of evolution - called "descent with modification" - was not
especially controversial, and several other evolutionary theories had already been proposed.
Darwin may stand at the beginning of a modern tradition, but he is also the final culmination of an
ancient speculation.
Greek Evolution
Many examples of societies that postulated the history of evolution include the Greeks, who did
not specifically refer to their concepts as "evolution", they did have a philosophical notion of
descent with modification. Several different Greek philosophers subscribed to a concept of
origination, arguing that all things originated from water or air. Another common concept was the
idea that all things descended from one central, guiding principle. Aristotle suggests a transition
between the living and the nonliving, and theorizes that in all things there is a constant desire to
move from the lower to the higher, finally becoming the divine.
Medieval Evolution
Medieval theories argued that all living things came into existence in unchanging forms due to
divine will, was notably in opposition to the concept of evolution. Medieval thinking was also,
oddly enough, confused by the idea of spontaneous generation, which stated that living things can
appear fully formed from inorganic matter. In this view, maggots came from rotting meat; frogs
came from slime, etc. This sort of a concept prevented both genetic thinking and speculation about
evolution or descent with modification. Nevertheless, a few philosophers theorized about some
sort of teleological principle by which species might derive from a divine form.
Immanuel Kant
Immanuel Kant the German philosopher Immanuel Kant developed a concept of descent that is
relatively close to modern thinking; he did in a way anticipate Darwinian thinking. Based on
similarities between organisms, Kant speculated that they may have come from a single ancestral
source. In a thoroughly modern speculation, he mused that "an orang-utan or a chimpanzee may
develop the organs which serve for walking, grasping objects, and speakingin short, that lie may
evolve the structure of man, with an organ for the use of reason, which shall gradually develop
itself by social culture".
The Ideas
Charles Darwin (1809-1882) is the father of evolution as we know it today (Campbell et al., 1999a;
Horan, 2006). His book “The origin of species” is the basis for modern evolution and speciation
theories. Darwin proposed that new species originate from ancestral species that change over time.
He added the mechanism of evolutionary change – natural selection (Freeman and Herron, 2004;
Grimaldi and Engel, 2005). Alfred R. Wallace (1823-1913) had ideas independent of Darwin and
was co-author with Darwin on the original paper proposing natural selection as the mechanism
behind evolution (Freeman and Herron, 2004; Grimaldi and Engel, 2005).
Darwin’s theory is based on four postulates from the introduction to “The Origin of species”
(Darwin, 1859). Darwin regarded life in nature as a competition, where the fittest individuals win.
The fitness of an individual refers to how well it survives and reproduces compared to other
individuals in the population. Traits that increase the fitness of an organism relative to individuals
without those traits makes it better adapted (Campbell et al., 1999a; Freeman and Herron, 2004).
However, Darwin could not explain how variation was passed from generation to generation and
how it was generated. That was solved by Gregor Mendel (1822-1884) and Thomas Hunt Morgan’s
(1866-1945) group at Columbia University. Based on experiments with pea plants, Mendel formed
laws about segregation and assortment of traits (Freeman and Herron, 2004; Grimaldi and Engel,
2005) and Morgan’s group demonstrated that Mendel’s hypothetical factors are specific points on
the chromosome (Allen, 1985a; Allen, 1985b).
In the 1920s Darwinian selection and Mendelian inheritance were integrated into the Modern
Evolutionary Synthesis by Dobzhansky, Mayr, Simpson and Stebbins (Campbell et al., 1999b;
Grimaldi and Engel, 2005; Freeman and Herron, 2004; Table 1). The synthesis emphasizes the
importance of populations as the unit of evolution; it states that mutations are the source of raw
material for evolutionary change, that natural selection is the most important mechanism of
evolution, and that large changes can evolve as the accumulation of small changes occurring over
long periods.
The classical Darwinian theories and the Modern Synthesis are now challenged, but these theories
have shaped most current ideas about evolution.
Level of selection
A central evolutionary concern is what unit is actually selected (Campbell et al., 1999a; Dawkins,
1976; Jablonka and Lamb, 2006). There are two general ways of looking at natural selection: the
gene's angle and that of the individual. The classical way is focusing on the individual, but Dawkins
(1976) introduced the gene's view of nature. Dawkins argued that genes and not the whole
organism is the unit of natural selection. The organisms are just “survival machines” for the genes.
However, the strictly gene-centred concept of natural selection may also be too simplistic, and the
two ways are probably equivalent (Freeman and Herron, 2004; Jablonka and Lamb, 2006). Natural
selection acts on phenotypes, but for evolution to occur there must be genetic variation that natural
selection can act on.
More than genes?
The phenotype of an individual is traditionally regarded as the summation of two totally
independent factors: the genes and the environment (Jablonka and Lamb, 2006). However, after
the introduction of epigenetic variation the separation is less clear. Epigenetic variation is inherited
variation that is sensitive to environmental input (Jablonka and Lamb, 2006; Lindqvist et al., 2007;
Richards, 2006). Traditionally, it was believed that inherited information only changes at random
and without direction towards a particular phenotypic outcome, but recent 14 findings indicate that
the environment can affect the genotype (Hoy, 2003). The most well known example is
transposable elements. Transposable elements are elements (with an RNA or DNA intermediate)
that can move from site to site in the genome (Hoy, 2003). The activity of the elements can be
induced by environmental factors, especially stress (Capy et al., 2000). This suggests that
transposable elements can create new genetic variation that is useful under conditions where the
fitness of an organism is reduced (Capy et al., 2000; Hoy, 2003). The evolutionary significance of
epigenetic mechanisms was first discovered in plants where the adaptive significance is clear
(Jablonka and Lamb, 2006). Plants cannot avoid harsh conditions by moving away and epigenetics
might allow them to respond in another way (Jablonka and Lamb, 2006).
Speciation
Despite the title of his book, Darwin devoted little space to the origin of species (Campbell et al.,
1999a; Coyne, 1994). He concentrated on how populations become adapted to their environment
through natural selection, but not how this adaptation leads to speciation (Campbell et al., 1999a;
Coyne, 1994). Now, the study of speciation is one of the most active areas of evolutionary biology,
and progress has been made in documenting and understanding phenomena in all aspects of
speciation (Turelli et al., 2001). However, there is a fundamental problem in the field. It is very
difficult to define exactly what “species” is. “It is as if on one hand we know just what “species”
means, and on the other hand, we have no idea what it means” (Hey, 2001). The idea of organic
discontinuity has a long tradition, beginning with Linnaeus’ classification (Coyne, 1994). The
clustering of organisms into discrete groups (i.e. species) can be seen in morphology, gene
sequences and reproductive compatibility (Turelli et al., 2001). However, some biologists argue
that the discontinuities are artefacts of human perception (Coyne, 1994), and in some groups e.g.
in plants and asexually reproductive taxa, it is difficult to separate different species (Coyne, 1994;
Turelli et al., 2001).
But why will organisms cluster into groups separated by gaps? And what properties of sexually
reproducing organisms and their environment lead to the evolution of discrete species? Two (not
mutually exclusive) explanations exist: the “ecological explanation” and the “sexual isolation
explanation” (Coyne and Orr, 2004; Turelli et al., 2001). The ecological explanation states that
ecological niches are discrete and that the clusters result from the ways different species exploit
physical resources. Furthermore, disruptive selection (Figure 1D) makes hybrids that “fall between
niches" less fit. The explanation for sexual isolation states that groups will adapt different to the
environment. Over time the number of differences will increase (divergent evolution) and result in
the formation of new species (Coyne and Orr, 2004).

Figure 1. Three general modes of selection. A) The original population. B) Stabilizing Selection:
Intermediate traits are favoured by selection, resulting in a decrease in variation. C) Directional
Selection: One extreme trait is favoured, resulting in a change in the mean value of the trait. D)
Disruptive Selection: Extreme traits are favored over the intermediate trait values, can divide the
population into two distinct groups. Disruptive selection plays an important role in speciation
(http://www.sparknotes.com/).
Species concepts
The biological species concept is classical and widely accepted (Berlocher, 1998; Campbell et al.,
1999b). It defines a species as a group of actually or potentially interbreeding populations that are
reproductively isolated from other such groups (i.e. they have the same gene pool). New species
arise when the evolution of reproductive isolation mechanisms prevents gene exchange between
populations (Turelli et al., 2001, Campbell et al., 1999c). Population biologists are discovering
more and more cases where the biological species concept is not valid e.g. in asexual organism
where the concept of breeding does not make sense. That results in the development of several
other species concepts (Box 1) (Campbell et al., 1999a; Coyne, 1994; Coyne and Orr, 2004;
Grimaldi and Engel, 2005).
Box 1. The biological species concept and some proposed alternatives (Campbell et al., 1999a;
Coyne, 1994; Coyne and Orr, 2004).

Biological species concept: Emphasizes reproductive isolation. Species are groups of actually
or potentially interbreeding natural populations that are reproductively isolated from other such
groups
Cohesion species concept: Focuses on mechanisms that maintain species as discrete
phenotypic entities. Each species is defined by its complex of genes and set of adaptations.
Applicable to organisms that reproduce without sex
Ecological species concept: Defines species on the basis of where they live and what they do
Evolutionary species concept: A species is a single lineage of ancestral and descendant
populations that are evolving independently of other such groups.
Genotypic cluster species concept: A species is a (morphologically or genetically)
distinguishable group of individuals that has few or no intermediates when in contact with other
such clusters
Morphological species concept: Defined species by measurable anatomical differences
(morphological criteria). It is practical to apply in the field, even to fossils.
Phylogenetic species concept: A species is the smallest monophyletic group of common
ancestry
Isolation of populations
Recognition species concept: Emphasizes mating adaption’s that become fixed in a population
as individuals “recognize” certain characteristics of suitable mates
Speciation in sexually reproductive organisms is based on the evolution of reproductive barriers
for the gene flow between populations (Campbell et al., 1999b; Turelli et al., 2001). Barriers can
occur before mating, between mating and fertilization, or after fertilization (Figure 2). Prezygotic
barriers occur before fertilization (figure 2) (Campbell et al., 1999b; Coyne and Orr, 2004). A
common prezygotic barrier is habitat isolation, where a geographical barrier (e.g. flooding) can
divide a population into several isolated populations (Campbell et al., 1999b).
Postzygotic barriers exercise isolation after fertilization (Figure 2; Table 1). The isolation can be
divided into extrinsic postzygotic and instrinsic postzygotic (Campbell et al., 1999b; Coyne and
Orr, 2004; Turelli et al., 2001). In extrinsic postzygotic isolation, hybrids are unfit because they
“fall between” parental niches as they have an intermediate phenotype that is less fit (Coyne and
Orr, 2004). In intrinsic postzygotic isolation, hybrids suffer developmental defects that make them
unable to survive or develop normally (Coyne and Orr, 2004).
Figure 2. The reproductive barriers that prevent gene flow between two different species.
Prezygotic barriers occurs before mating, while postzygotic do after (Campbell et al., 1999b).
Table 1. Classification of postzygotic reproductive isolation (Coyne and Orr, 2004)
Types of speciation
There are two general modes of speciation: allopatric speciation and sympatric speciation
(Figure 3). They are defined by how the gene flow among populations is interrupted. In allopatric
speciation a geographical barrier physically isolates a population and initially blocks gene flow,
whereas in sympatric speciation intrinsic factors e.g. chromosomal changes or nonrandom mating
alter the gene flow.

Figure 3. The two general modes of speciation. Top) allopatric speciation. Bottom) sympatric
speciation.
Allopatric speciation
In allopatric speciation populations are separated by geographical isolation. In allopatric speciation
extrinsic factors – as great distance or a physical barrier – prevents two or more groups from mating
(Campbell et al., 1999b). Physical isolation is an effective barrier to gene flow and in many cases
it is an important trigger for divergence. When no forces impose reproductive capability between
isolated populations the populations will, given enough time, become incompatible (Turelli et al.,
2001). Allopatric speciation is most likely to occur if a small population in the periphery of a
species’ range gets isolated. The individuals in the periphery are often extremes with a gene pool
that differs from that of the rest of the population (Campbell et al., 1999b; Freeman and Herron,
2004). In a small population random mutations or new combinations of existing alleles with neutral
adaptive value may get fixed by chance and evolution by natural selection may be different than
in the parent population (Campbell et al., 1999b).
Sympatric speciation
Since the nineteenth century it has been debated if speciation requires geographical isolation
(Berlocher, 1998). The authorities (e.g. Mayr and Dobzhansky) argued that geographic isolation is
a necessary first step for divergence in animals whereas Guy Bush emphasized ecological adaption
as an important factor in speciation (Bush, 1998; Feder et al., 2005). Sympatric speciation is still
questioned and recent analyses show that allopatric speciation is the most common mode
(Barraclough and Nee, 2001).
Two central factors differ between sympatric and allopatric speciation. Firstly, sympatric
speciation does not require large-scale geographic distance to reduce gene flow between parts of a
population (Campbell et al., 1999b; Freeman and Herron, 2004). Instead new species arise within
the range of the parent population as the result of reproductive barriers between the mutant and the
parent populations. Secondly, in sympatric speciation gene flow may continue for a number of
generations after the populations have become separated, whereas complete isolation arises
between populations evolving in allopatry.
A four stage series has been proposed for sympatric speciation via host plant shift for phytophagous
insects (Berlocher, 1998): (1) partially reproductively isolated host races (2) species isolated only
by host fidelity (3) species with partial prezygotic and/or postzygotic isolation unrelated to host
fidelity and (4) totally reproductively isolated species.