BIOL2007 TUTORIAL EXERCISE:

Due tomorrow, Fri. 4:30

Kevin in office 2:30-4:00 pm
TODAY
May be some help if you are having
problems with game theory exercise
BIOL2007 EVOLUTION OF SEX AND
SEXUAL SELECTION

EVOLUTIONARY QUESTIONS
ABOUT SEX
Today:
A) The evolution of sex
1) What is the advantage of sex?
 
B) Ev. of sexual dimorphism - sexual selection
2) Why are there "secondary sexual
characteristics"
OTHER interesting questions could be asked:

4) Why meiosis?
5) Why mitochondria &c asexual?
6) Why only 2 sexes?
7) What is the optimal sex ratio? (Kevin)
8) Why does the Y chromosome become
degenerate (non-coding)?
9) How does sex determination evolve?
10) Sexual conflict – why and how?
 
See: BIOL2011 (Behavioural Ecology & Sociobiol)
and BIOL3012 (Sex, Genes and Evolution).
WHAT IS SEX?
 Recombination – all of life
 Meiosis - eukaryotes only
 Anisogamy - unequal gamete size - multicell eukaryotes
 Dioecy - Separate sexes
- vs. hermaphrodites and monoecy
 
These are complex energy-requiring traits
 ADAPTATIONS

Recombination seems to be a lowest common denominator

of all sex.
EVOLUTION OF SEX

Costs of sex
 
Recombination and sex is complicated,
costly, predation-prone (even if fun).
Two-fold cost of sex
Populations grow faster if parthenogens.
Suppose each female produces 2.4
progeny:

... and so on (but it works for any reprod. rate)

Males contribute little; males are useless!

SO: parthenogens should outdo sexual

females and increase in frequency.

(But advantage not always twofold…)

 
If such strong disadvantages to dioecy,
…must be some pretty hefty advantages.

Recombination is the primary feature of sex.

Advantages of sex must have something to
do with recombination.
Advantages of sex

a) recombination increases evolutionary rate

Muller: Recombination allows advantageous mutations
to combine in the same individuals.
 D'you know what dandelions
Higher evolutionary rate can be adaptive for a species.

May work against individual are?

disadvantages for sex within
populations: so group selection.

Asexual "cheats" gain individual advantage, spread to

fixation within species, not to other species.

Asexual species do exist (e.g. dandelions). Why do they

not take over?

If other species evolving/adapting faster, competition

may cause extinction of asexuals  one of the few cases
where group selection may actually be operating.

(Conversely, why don't dandelions die out?)

Evidence:
asexual species trapped on “twigs” in phylogenies.
b) Individual selection
Survival in a coevolutionary "arms race“

Environment of offspring may be very different from

that of their parent, parent’s adaptations may not be
sufficient (e.g. Daphnia, aphids).

Sex as a lottery: at least some progeny with

reshuffled adaptations - direct individual selection
advantage to parents in producing recombined,
variable offspring.

Other ideas. There are plenty, but we will leave that

to other courses.
EVOLUTION OF SEXUAL DIMORPHISM -
SEXUAL SELECTION

Darwin: secondary sexual characteristics due to a

struggle for genetic representation between
members of the same sex.

Sexual selection (struggle for mating success) vs.

natural selection (struggle for survival and
reproduction after mating).

Nowadays: sexual selection a special form of

natural selection, but a very important one.
Birds of
paradise
(Paradiseidae)
Western grebe
Aechmophorus
occidentalis
Jackson’s widowbird
Euplectes jacksoni
Satin bowerbird
Ptilinoorhynchus
violaceus
Stag beetle
Lucanus cervus
Stalk-eyed fly
Cyrtidopsis
dalmanii
(Diopsidae)
Why are males more extreme?

Males usually more flamboyant than females,

with horns, bright colours, or displays.

Why?  Darwin's view: males are more

"passionate" than females. So females can be
more choosy. Males have to “sedulously
display their charms before the females”

Maybe (!); but why are males more

passionate?
Males:
cheap gametes
can produce lots
little parental investment
 potentially father many offspring

Females:
expensive gametes
produce few
often show more parental care,
at least as far as nutrients in the egg
 more limited no. offspring than males
 Males:
compete for access to females
more indiscriminate, less to lose

 Females:
plenty of willing males around
worthwhile to be choosy, because number
of matings doesn’t limit offspring number
Elephant seals:
> 90% males father no
offspring;
fittest fathered 93.
But > 50% females have
1 offspring. 

Males: higher variance in

offspring number.
Sexual selection can lead to increased matings
at the expense of reduced survival.

Darwin : sexual selection might outweigh

natural selection, and explain conspicuous
coloration in the animal kingdom.

e.g. sexual selection may antagonize natural

selection for camouflage.
Two major types of sexual selection:
Intrasexual selection - male-male
competition
in which males compete by fighting other
males for females

And …

Intersexual selection - sexual selection by

female choice
in which males compete to obtain the most
interest from females.
Intrasexual selection - male-male competition
a) Selection for fighting ability
Examples:

 Red deer males: harems of females, defend by

roaring, displaying, and finally fighting; injuries and
death may result

Salmon: males fight to the death with enlarged

hooklike jaws, useless for feeding.

 Fig wasps: specialized males with huge jaws hatch

out and kill other males, then mate with all other
females in the fig.
b) Sperm competition

Male-male competition: after mating.

Sexual selection between sperm of rival males for
fertilization in reproductive tract of females.

In primates, males from species with

polyandry have larger testes than males from
monandrous species.  e.g. Gorillas
monandrous, chimps polyandrous.

See also: other courses

RA Fisher:
his analyses rehabilitated female choice

Today, explosion of work showing:

1) females do indeed choose, and

2) sexually selected traits often costly.

 Before Widowbird
Elongated Euplectes
Control I progne
Control II
Shortened

0.0 0.5 1.0 1.5 2.0

After

Elongated
Control I
Control II
Shortened

0.0 0.5 1.0 1.5 2.0

Average no. mates
Why SHOULD females choose males with
exaggerated and costly traits?
Three major theories:

a) Sensory bias. Exaggeration is a

supernormal sign stimulus.

Pre-existing preferences: Physalaemus frog

phylogeny shows that "chuck" sound has
evolved in a terminal branch of the genus.

Females of related spp. which lack "chuck"

still like it. (Mike Ryan).
a) Sensory bias

b) Direct benefits of female choice

e.g. avoiding disease by mating with
uninfected males. Or getting help at
the nest

c) Indirect benefits of female choice

Indirect benefits are passed on to the
offspring via genetic contribution
from male.
i) “sexy sons”
Mathematical models, by Russ Lande
and Mark Kirkpatrick (1980s): runaway
models of sexual selection.
Imagine male trait evolution and female choice; females can
choose whatever they want; males have a natural-selection
optimum.
Coevolution of male trait and female response  potential
runaway process of evolution.
ii) "good genes" hypothesis
Exaggerated, costly traits  high fitness.
 
Parasite hypothesis of Hamilton and Zuk
Male birds and mammals: bare patches of skin
(e.g. baboons’ bottoms, wattles on cockerels)

Diseases, parasites: alters blood colour

 Bare flesh: honest signals of overall health.

Hamilton & Zuk: sexually dimorphic species of

birds suffer more from parasites.
Zahavi’s "handicap principle".
Not very costly male badges prone to
cheating

Zahavi: only very costly traits uncheatable

A costly "handicap" is selected, paradoxically,

because it guarantees "honest signalling".
Which is correct?
Direct selection? Clearly this occurs

"Good genes" some evidence, easy to test

Non-adaptive "sexy sons" idea appealing, but

difficult to prove

Evolutionists haven’t yet resolved these

arguments, but they are fun to think about
Sexual selection in humans

Darwin: "The Descent of Man and

Selection in Relation to Sex".

Sexual selection may explain the

extraordinary racial differences in
animals and humans
Desmond Morris, Jared Diamond,
“evolutionary psychologists”:
similar conclusions

Peculiarly human features such as:

• bare skin, hairlessness
• lips
• hair colour, skin colour
• enlarged breasts in females
• beard in males
• copulation when not in oestrus
• weapons, war?
• ornaments, jewellery?
• music?
• art? ... etc.
TAKE HOME POINTS

Evolution of sex:
• The basis of sex is recombination
• Sex is usually costly
• Separate sexes may exist because of:
 asexual lineages die out (group selection)
 offspring variability (individual selection)
TAKE HOME POINTS (contd.)

Sexual selection/sexual dimorphism:

• Male-male competition
 fighting
 sperm competition

• Female choice
 sensory bias
 direct benefit to self/offspring
 indirect (inherited benefits)
 sexy sons
 good genes
FURTHER READING

FUTUYMA, Evolution.
Ch 14: 329-339, Ch 17: 417-426.

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