ASP1022 Topic 13

Evolution and Natural Selection

Natural selection: A not-so-simple idea
Charles Darwin was not the rst European scientist to believe
that species change, and that new species arise over time.
The 'transmutation of species' (to use the 19th Century term)
had been a commonly accepted idea among scientists for 50
years before Darwin published his great work On the Origin of
Species by means of Natural Selection in 1859. The
mechanism driving the change, however, was unknown until
Darwin proposed that natural selection was responsible. The
idea was immediately hailed by the scientic community, and
remains today as the
cornerstone of
evolutionary theory.
The basic idea of natural selection can be stated in everyday language:
1. Individuals in any species can potentially reproduce at a rate which would quickly ll the world with sea
urchins, oak trees, elephants, or whatever species we want to consider.
2. Since this doesn't happen, the full potential fecundity
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of individuals must be thwarted: not all individuals
survive, and not all survivors reproduce.
3. Which individuals survive, and which reproduce, is determined by competition among individuals. There is
variation among the individuals of any species, so those with traits which make them superior competitors
will be the most successful at survival and reproduction.
4. Provided that these 'winning' traits have a genetic basis (Darwin and other 19th Century scientists knew
next-to-nothing about genetics!), they will be passed on to offspring and will become more common in future
generations.
5. Genetic mutations (which occur every time a cell divides, due to rare errors made by the DNA-copying
mechanism) continually supply new variation on which natural selection can act. This process can, over long
periods of time, result in new species.
Perhaps because this core argument seems simple, people have used little more than the catchphrase
'survival of the ttest' to make arguments about how evolution works, and what (or whom) natural selection
favours. A central goal of this lecture is to convince you that natural selection isn't so simple: it often acts in
subtle and unexpected ways.
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fecundity = ability to reproduce abundantly
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Before
reading further, briey note
down what you think is meant
by 'natural selection'
Quick quiz! (answer before you read the next page!)
Can natural selection favour early death?
Does natural selection affect the sex-ratio of a species? If so, how?
Does natural selection lead to stronger species?
A surprise: natural selection can favour genes that cause early death
At rst, this idea seems completely contrary to evolutionary theory: isn't natural selection about survival of
the ttest, not early death of the ttest?
The important point to realise here is that survival alone is irrelevant. Survival is important to evolutionary
tness only if it eventually enhances reproduction.
In many species, including humans, genes that affect ageing (including genes which affect susceptibility to
diseases which typically strike in old age) are known to have 'trade offs' with reproductive success earlier in
life. These genes promote fecundity at young ages, but impose a cost to survival later in life.
For example, experiments with the fruit y Drosophila melanogaster, a favourite lab organism of geneticists,
have shown how articially imposed selection for longer life span can work: the resulting ies live almost
twice as long as ies in a control group, but they are less successful at reproducing at earlier ages.
Indeed, ageing and death in many (perhaps all) organisms seems to be due (at least in part) to the fact that
biochemical and physiological maintenance of the body isn't worth the cost of lower reproductive success. In
natural selection terms, it is better to die young if doing so enhances your overall reproduction.
Another surprise: selection and the sex ratio
Most species have a 1:1 sex ratio of males to females. Why is this?
One easy answer (for anyone who knows some basic genetics) is that the X and Y sex-determining
chromosomes ensure a 1:1 ratio. This seems like a good idea, but can't explain everything that is seen; for
example, some organisms, such as crocodiles and turtles, lack sex chromosomes entirely, yet they still have
a 1:1 sex ratio!
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So there must be more to it
Another initially persuasive, but incorrect, argument is based on the observation that males in many species
ght for control of a harem or mating territory. The winners in this contest might have several mates, while
the losers could never reproduce in their lifetimes. It is tempting to suppose that natural selection produces a
1:1 ratio to promote this competition for mates, so as to ensure that only the most genetically superior males
pass genes to the next generation. This hypothesis is also incorrect.
Natural selection is not concerned about the 'health', 'vigour' or 'genetic tness' of any species; it is merely a
blind mechanism by which those who reproduce the most pass on their traits to future generations. Natural
selection does not have goals, such as keeping a species t. If natural selection simply brought about 'good
outcomes', and it it were good for the species to have males compete for mates, why should the sex ratio be
1:1? Why not favour 60% males, or 90%, or 99%? Then there would be lots of competition. Competition
among males for mates is not the cause of the 1:1 sex ratio; it is a consequence.
The true role of natural selection involves a process called frequency-dependent selection, in which average
tness is greater for the minority sex.
Suppose a population had a female-biased sex ratio, say three females to every male. As a parent, would
you be better off producing sons or daughters?
For every daughter you produce, you have on average, the grand-offspring that one female can produce. But
because males can, on average, obtain three mates each (3:1 sex ratio), every son you produce will give
you, on average, the grand-offspring of three females. Thus you are better adapted ('more t') in the long run
if you produce sons.
(Of course, it isn't up to an individual's voluntary desire. Rather, any genetic mutation which tends to increase
the production of sons over daughters would be favoured by frequency-dependent selection in this instance.
Biologists have identied many genes that have such effects in many organisms).
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In these species, the gender of an individual is determined by the temperature it experiences during incubation of its
egg. A 1:1 sex ratio is achieved because the genes that set the temperature threshold between male and female are
'calibrated' by natural selection so that approximately equal numbers of each sex are produced, given the average range
of temperatures experienced by eggs of that species.
In contrast, if the sex ratio is biased 3:1 in favour of males, and you produce sons, most on average will
never mate. The average reproductive success of a son would be 1/3 that of a daughter, in which case
natural selection will favour those traits which produce female offspring.
The equilibrium point of these opposing actions of frequency-dependent selection is at a 1:1 ratio. This is the
fundamental reason for equal numbers of each sex. (The XY chromosome system has evolved in some
species as a mechanism to bring about a 1:1 ratio, but it is not the fundamental reason behind it).
Natural Selection and Improvement
The phrase 'survival of the ttest' might also seem to imply that species must continually improve over time.
Al elegant experiment conducted with yeast shows that this is not really the case.
Yeasts are single-celled organisms which absorb sugars from their environment. A group of researchers
cultured yeast populations in solutions with very low sugar concentration. Any mutation which improves a
cell's ability to compete for the limited sugar would allow hat mutant lineage to grow faster than its
competitors (remember that mutations occur every time a cell divide, due to the less-than-100% accuracy of
the copying mechanism. In a yeast culture of millions of cells which divide every few hours, billions of
mutations will occur in a relatively short time).
Using molecular biology techniques, the researchers determined that ve mutant lineages arose in
succession over the course of 300 generations of yeast cells. Each new mutant lineage out-competed its
predecessor lineage for the available sugar, and eventually completely replaced the preceding lineage, until
it too was later replaced by a superior successor.
One might suppose that the 5th lineage to appear must be a better competitor than all the previous lineages.
After all, lineage 5 beat lineage 4, which beat lineage 3, which out-competed lineage 2, and so on. But, this
was not the case. When an extract of lineage 5 was allowed to compete with yeast cells from lineage 2, the
lineage 2 cells grew 10% faster than those from lineage 5. Similarly, the original lineage of cells was 3%
better than lineage 4 when both were placed in competition with each other.
This clever experiment reinforces the point made above: natural selection does not have a goal of making
species better. It is simply a mechanism which favours the spread of traits that allow superior survival and
reproduction relative to the existing population.
Conclusion
Natural selection is a complex, and often counter-intuitive process. It is a mechanism which operates on
random mutations. It does not have a direction, or seek to make organisms better over time. Instead it just
works to select traits within a population which are best adapted to the current environment and conditions. It
will not result in perfection; just better adaptation.
The main driving force for natural selection is enhanced reproductive success.
Points to ponder
Does natural selection work on species or individuals?
Does natural selection have a goal?
Why is the term 'survival of the ttest' an over-simplication of the process?
Why do many species have a 1:1 sex ratio?
Are humans subject to natural selection?
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