Aging and Other Life-History Characters

Basic issues in life-history

o Reproductive success is the trait on which natural selection acts
o Life history attempts to make sense of the diversity in
reproductive strategies
o Organisms face fundamental trade-offs in their use of energy and
time
Amount of energy an organism can harvest is limited
Energy and time tradeoff
Size tradeoff energy cost
Time required to grow is time during which predators,
diseases, or accidents may strike
An individual that takes the time to grow large thus
incurs a greater risk of dying without reproducing
Whenever there is a trade-off between different
components of fitness, we expect natural selection to
favor individuals that allocate energy and time with
an optimal balance between benefits and costs,
thereby maximizing lifetime reproductive success
o Changes in life history are caused by changes in allocation of
energy
Allocating less energy to reproductions means having
smaller litters
Natural selection on life histories leads to adjustments in
energy allocation that maximize the total lifetime
production of offspring
Why do organisms age and die?
o Aging (senescence) is a late-life decline in an individuals fertility
and prob- ability of survival
o Aging reduces an individuals fitness
Aging should be opposed by natural selection
o Rate of living theory
Populations lack the genetic variation to respond any
further to selection against aging
Aging caused by accumulation of irreparable damage to
cells and tissue
Damage caused by errors during replication
Accumulation of metabolic products
All organisms have been selected to resist and repair cell
and tissue damage to the maximum extent physiologically
possible
Have reached the limit of biologically possible repair

Populations lack the genetic variation that would

enable them to evolve more effective repair
mechanisms that they already have
Aging rate should be correlated with metabolic rate
Species should not be able to evolve longer life spans
whether subjected to natural or artificial selection
According to the theory, all should expend about the same
amount of energy per gram per lifetime, whether they burn
it slowly over a long life or rapidly over a short one
BUT data on variation in metabolic rate and aging
among mammals refute this theory
Organisms age and die due to intrinsic physiological limits
on cells and tissues (little flawed)
o Evolutionary theory
Trade-off between allocation of energy to reproduction
versus repair
Aging is caused not so much by cell and tissue damage
itself as by the failure of organisms to completely repair
such damage
This failure leads to gradual decay and ultimate
collapse
Failure to completely repair damage is ultimately caused
by either deleterious mutations or trade-offs between
repair and reproduction
Natural selection is weak late in life so alleles that cause
aging are only mildly deleterious
They may persist in mutation-selection balance or
rise to high frequency by drift
In populations where mortality rates are high, individuals
tend to invest more heavily in early reproduction
The evolutionary theory of senescence hinges on the
observation that the power of natural selection declines
late in life
This is because most individuals diedue to
predators, diseases, or accidentsbefore reaching
late life.
Two mechanisms can lead to the evolution of
senescence:
o (1) Deleterious mutations whose effects occur
late in life can accumulate in populations;
o and (2) when there are trade-offs between
reproduction and maintenance, selection may
favor investing in early reproduction even at

How
o
o

the expense of maintaining cells and tissues in

good repair
Evolutionary explanation for menopause
In humans, reproductive capacity declines earlier and more
rapidly in women than men
Nonadaptive artifact of our modern lifestyle
Reconstructing the demography of ancient peoples
have often concluded that in premodern cultures,
virtually all adults died by age 50 or 55
Not an adaption because hunter-gatherer ancestors
never lived long enough to experience it
Life History adaptation
Menopause- life history adaption associated with the
contribution grandma makes to feeding her
grandchildren
Human children depend on mom for food several
years after weaning
1. Probability that she will live long enough to be able
to nurture another baby from birth to independence
declines
2. risks associated with pregnancy and childbirth rise
3. her own daughters will themselves start to have
children
suggests that older women may reach a point at which
they can get more additional copies of their genes into
future generations by ceasing to reproduce themselves and
instead helping to provision their weaned grandchildren so
their daughters can have more babies.
grand- mothers face a trade-off between investment
in children versus grandchildren
age at menopause is heritable and later menopause
confers higher lifetime reproductive success
many offspring should an individual produce in a given year?
Trade offs: more offspring a parent (or pair of parents) attempts
to raise at once, the less time and energy the parent can devote
to caring for each one
Clutch size in birds
Selection will favor clutch size that produces the most
surviving offspring
Probability that any individual offspring will survive
decreases with increasing clutch size
Adding eggs reduces survival rate for individual
chicks

o Ability of parents to feed any individual

offspring declines as the number of offspring
increases
Number of surviving offspring reaches a maximum at an
intermediate clutch size. It is this most-productive clutch
size that Lacks hypothesis predicts will evolve by natural
selection
Lacks hypothesis predicts that parents will attempt to rear
that number of young that maximizes the number of
surviving offspring.
Data indicate that parents often rear fewer offspring.
Efforts to identify which of Lacks assumptions are
violated have led to the discovery of additional tradeoffs and improved estimates of lifetime fitness
o Criticisms of Lacks hypothesis
1. Lacks hypothesis assumes there is no trade-off between
a parents reproductive effort in one year and its survival or
reproductive performance in future years
When reproduction is costly and selection favors
withholding some reproductive effort for the future,
the optimal clutch size may be less than the most
productive clutch size
2. Lacks hypothesis assumes that the only effect of clutch
size on off- spring is in determining whether the offspring
survive. Being part of a large clutch may, however, impose
other costs on individual offspring than just reducing their
probability of survival
clutch size affects not only offspring survival but also
offspring reproductive performance
When larger clutches entail lower offspring
reproductive success, the optimal clutch size will be
smaller than the most numerically productive clutch
size
3. discrepancy between Lacks hypothesis and the behavior
of individual birds may sometimes be more apparent than
real
Note: assumed that clutch size is fixed for any given
genotype. In fact, clutch size is often phenotypically
plastic
o If clutch size is plastic, and if birds can predict
whether they are going to have a good year or
a bad year, we would predict that individuals
will adjust their clutch size to the optimum
value for each kind of year
o Larger clutch sizes may reduce offspring fitness

Trade-offs between a females investment in a particular

clutch and her own future survival or reproductive
performance
big should each offspring be?
Individual offspring will have a better chance of surviving if they
are larger
Minimum size below which offspring have no chance of
survival. As offspring get larger, their probability of
surviving rises
Expected fitness of a parent producing offspring of a
particular size is the number of such offspring the parent
can make multiplied by the probability that any individual
offspring will survive
Key point: selection on parents often favors offspring
smaller than the size favored by selection on offspring
themselves
Survival probability increases with offspring size, but the
rate of increase decline
Increasingly large offspring gain a progressively
smaller survival benefit
Intermediate offspring size give the highest parental
fitness
1. Trade off between size and number of offspring
2. Above minimum size, probability that any individual offspring
will survive is an increasing function of its size
Problem: organisms face a trade-off between making many lowquality offspring or a few high-quality offspring. Selection on
parents favors a compromise between the quality and quan- tity
of offspring, but selection on individual offspring favors high
quality.

How
o

o
o
o