THE UNIT OF

SELECTION
Part 3 Adaptation and Natural Selection
Fathan Hadyan R
3415106786

Rahmat Fadrikal
3415096616
What entities benefit from the
adaptations produced by
selection?
It is a familiar idea that life can be divided into
a series of levels of organization : nucleotide
to gene, through cell, organ, and organism, to
social group, species, and higher levels
In some cases, what benefits an organism
may not also benefit its species,
in these cases the evolutionary biologist
needs to know which level natural selection
most directly benefits
If we are seeking to understand why an
adaptation evolved, we need to know what
entities adaptations in general evolve for the
benefit of.
Lions often hunt alone, but they can improve their
chance of success by hunting in a group

When prey have been detected, a wildebeest herd
perhaps, the lions start to stalk towards them. As
they get close, they take different routes, some
going on straight ahead, and some to the sides, so
the prey herd is approached by lions stalking them
from different directions. . . . Eventually one lion
gets close enough to make a rush at a wildebeest,
or else a lion is detected by the prey Bertram
(1978).
The cooperative behavior of the hunting party is
here an adaptation for catching food, but it is not
the lions only feeding adaptation. The lions
muscular jaws and limbs, their teeth and five
senses, all contribute to the success of the hunt
Each time a hunt is successful, there will be a
small incremental increase in the species chance
of survival, or avoiding extinction.
The survival probability will also be increased, if
by a smaller amount, for the genus Felis and the
cat family Felidae.
The beneficial effect of the hunt spreads
downwards as well as up from the individual
lion
As the survival of the lion is increased, so is
the survival of its constituents ( the organs,
cells, proteins, and genes)
The levels of organization, from gene through
individual lion to Felidae, are to a large extent
bound together in their evolutionary fate, and
what benefits one level will usually also benefit
the others
However, this is not always so. Male lions can
only join a pride by forcibly evicting the
incumbent males. In the fight, lions may get
killed or wounded, and in any case lions have
a low rate of survival after they have been
evicted from a pride
The survival of the species may be little
affected by the death of male lions, because
the mating system is polygynous and has
plenty of males to spare; but the effect is
clearly not positive


the levels of living organization are bound
together, if natural selection produces an
adaptation to benefit one level, many other
levels will benefit as a consequence
if it benefits one level, which is it?
What is the unit of selection?
Natural selection has produced
adaptations that benefit various
levels of organization
Segregation distortion benefits one gene at
the expense of its allele
Selection may sometimes favor some cell
lines relative to other cell lines in the same
body
Natural selection has produced many
adaptations to benefit organisms
Natural selection has produced
adaptations that benefit various
levels of organization
Natural selection working on groups of close
genetic relatives is called kin selection
Whether group selection ever produces
adaptations for the benefit of groups has been
controversial, though most biologists now think it
is only a weak force in evolution
Which level in the hierarchy of organization levels
will evolve adaptations is controlled by which level
shows heritability

Segregation distortion benefits
one gene at the expense of its
allele

With normal Mendelian segregation at a
genetic locus, on average half of an
organisms offspring inherit one of the alleles
and the other half the other allele.
Mendelian segregation is so to speak fair in
its treatment of genes: genes emerge from
Mendelian segregation in the same
proportions as they went in
some curious cases in which Mendels laws
are broken in which one of the alleles, instead
of being inherited by 50% of a heterozygotes
offspring, is consistently overrepresented
The segregation distorter gene of Drosophila
melanogaster is an example of this
phenomenon, which is also called meiotic
drive.
A heterozygote for the segregation distorter is
then sd/+. The majority (90% or more) of offspring
from male heterozygotes have the sd gene
because the sperm containing the + gene fail to
develop. Female heterozygotes have normal
Mendelian segregation.
A segregation distorter gene can have a great
selective advantage. The allele that gets into more
than half of a heterozygotes offspring will
automatically increase in frequency and should
spread through the population.
Segregation distortion sets up an interesting
selection pressure in the rest of the genome.
On average, all other genes at other loci suffer
a disadvantage because of segregation
distortion.
When selection acts in conflicting ways on
different genes in the same individual body, it
is called intragenomic conflict. The sd/+ fruitfly
has intragenomic conflict, because selection
on the sd gene favors segregation distortion
and selection on other genes favors restoring
normal segregation.

Selection may sometimes favor
some cell lines relative to other
cell lines in the same body

In organisms like ourselves, a new individual
develops from an initial single-cell stage and
that single cell derives from a special cell line,
the germ line, in its parents
In a Weismannist organism, most cell lines
(the soma) inevitably die when the organism
dies; reproduction is concentrated in a
separate germ line of cells.
The separation of the germ line limits the
possibilities for selection at the suborganismic
level, between cell lines. One cell may mutate
and become able to out-reproduce other cell
lines and (like a cancer) proliferate through the
body.
Any somatic cell line comes to an end with the
organisms death. For this reason, cell
selection is not important in species like
ourselves.
However, Buss (1987) pointed out that
Weismannist development is relatively
exceptional among multicellular organisms
We tend to think of it as usual because
vertebrates, as well as the more familiar
invertebrates like arthropods, develop in a
Weismannist manner. However, more than half
the taxa listed in Table 11.1 have the capacity
for somatic embryogenesis : a new generation
may be formed from cells other than those in
specialized reproductive organs.
Tabel 11.1 s: somatic ; p:preformationistic ;
e:epgenetic ; u:unknown (developemental
mode)
In a species in which new offspring can
develop from more than one cell lineage,
selection between cell lines becomes possible.
When the organism is conceived it will be a
single cell, and for the first few rounds of cell
division the organism will probably remain
genetically uniform.
In a Weismannist species, that cell line will die
when the organism dies and any selection
between cell lines will be unimportant. However, if
any cell line in the body has some chance of
giving rise to the next organismal generation, the
mutant cell line would increase its chance of being
in an offspring and be favored by selection.
Whitham & Slobodchikoff (1981) argued that in
plants selection between cell lines enables the
individual to adapt to local conditions more rapidly
than would be possible with strictly Weismannist
inheritance
The process is at present more of a theoretical
possibility than a confirmed empirical fact, but
it may well be important in non-Weismannist
species
Natural selection has produced
many adaptations to benefit
organisms
the beaks of the woodpecker
the Galpagos finches
the peppered moth Biston betularia
mimetic butterflies

are all adaptations that benefit the individual
organism. It can hardly be doubted, therefore,
that organismal adaptations exist, and natural
selectioncan favor them.
Natural selection working on
groups of close genetic relatives is
called kin selection
In species in which individuals sometimes
meet one another, such as in social groups,
individuals may be able to influence each
others reproduction
Altruistic behavior often takes place between
genetic relatives, and when it does the most
likely explanation is the theory of kin selection.
Under what condition will natural selection
favor altruism?
The altruist still pays a cost of c for performing
the act, and the recipient receives a benefit b
However, the chance that the altruistic gene is
in the recipient is r. When rb exceeds c there
will be a net increase in the average fitness of
the altruists
The condition for natural selection to favor
altruism among relatives is that it should be
performed if :
rb > c
This is the theory of kin selection. It states that
an individual is selected to behave altruistically
provided that rb > c. The condition itself is
called Hamiltons rule
Whether group selection ever
produces adaptations for the
benefit of groups
A group adaptation is a property of a group of
organisms that benefits the survival and
reproduction of the group as a whole
group adaptations that did not evolve by kin
selection
Many characters are beneficial at the group
level, but also benefit all the individuals in the
group
The controversial group adaptations are those
that benefit the group but not the individual
A hypothetical example is Wynne-Edwards
theory, put forward in Animal Dispersion in
Relation to Social Behaviour (1962), that
animals restrain their reproduction in order not
to overeat the local food supply.
Many models of group and individual selection
exist, but they can mainly be reduced to a
common form (Figure 11.1).
The groups are supposed to occupy patches in
nature. As before, some patches are occupied by
altruistic and others by selfish groups. There are
also empty patches. A selfish group in the model
drives itself extinct by overeating its patchs
resources. The result of the model depends on
whether a selfish group can infect an empty or
altristic patch before going extinct.
Figure 11.1
Maynard Smiths
formulation of group
selection models. A
patch may change
state in the direction
of the arrows.
Redrawn, by
permission of the
publisher, from
Maynard Smith
(1976). 1976
University of Chicago
Press
Maynard Smith
(1976) defines the number m as the number of
successful migrants produced by one selfish
group on average between its origin and
extinction. (Successful means that the migrant
establishes itself in another group and breeds.) If
m= 1 the system will be stable;
if m < 1 the selfish groups decrease in number,
and if m > 1 they increase. In other words, a
selfish group only needs to produce more than
one successful emigrant during its existence for
the selfish trait to take over
Which level in the hierarchy of organization
levels will evolve adaptations is controlled by
which level shows heritability
Most adaptations appear to benefit the
individual organism
Heritability is the key to which levels of
organization show adaptations
Another sense of unit of selection is the
entity whose frequency is adjusted directly by
natural selection
The unit of selection, in a second sense, is the
gene, because only genes last long enough for
natural selection to adjust their frequencies.
The longevity of a genetic unit matters relative
to the time evolutionary change takes
The word gene is being used in a technical
sense
Genes are actively, not passively, part of
natural selection