Storage for Good Times and Bad:

Of Squirrels and Men

Test

A fable of food-hoarding,
As in Aesop and Walt Disney
The fable concerns squirrels, but has more
ambitious intentions.
What can evolution tell us about the evolution of
our preferences toward risk?
For the moral of the story, we look to the works of
another great fabulist
Art Robson

Preferences toward risk

Robson (JET 1996) : Evolutionary theory
predicts that:
For idiosyncratic risks, humans should seek
to maximize arithmetic mean reproductive
success. (Expected utility hypothesis.)
For aggregate risks, they should seek to
maximize geometric mean survival
probability.

A Simple Tale
Squirrels must gather nuts to survive
through winter.
Gathering nuts is costlypredation risk.
Squirrels dont know how long the winter
will be.
How do they decide how much to store?

Assumptions
There are two kinds of winters, long and
short.
Climate is cyclical; cycles of length
k=kS+kL, with kS short and k L long winters.
Two strategies, S and L. Store enough for a
long winter or a short winter.
Probability of surviving predators: vS for
Strategy S and vL=(1-h)vS for Strategy L.

Survival probabilities
A squirrel will survive and produce
offspring iff it is not eaten by predators and
it stores enough for the winter.
If winter is short, Strategy S squirrel
survives with probability vS and Strategy L
with probability vL<vS.
If winter is long, Strategy S squirrel dies,
Strategy L squirrel survives with prob vL

No Sex Please
Reproduction is asexual (see Disney and
Robson). Strategies are inherited from
parent.
Suppose pure strategies are the only
possibility.
Eventually all squirrels use Strategy L.
But what if long winters are very rare?

Can Mother Nature Do Better?

How about a gene that randomizes its
instructions.
Gene diversifies its portfolio and is
carried by some Strategy S and some
Strategy L squirrels.
In general, such a gene will outperform the
pure strategy genes.

Random Strategy
A randomizing gene tells its squirrel to use
Strategy L with probability L and Strategy
S with probability S.
The reproduction rate of this gene will be

SS()= vS S+vL L, if the winter is short.

SL()=vL L

if the winter is long.

Optimal Random Strategy

Expected number of offspring of a random strategist
over the course of a single cycle is
kSS() kSSL() kL
Optimal strategy chooses probability vector
=(L ,S ) to maximize above.
A gene that does this will reproduce more rapidly
over each cycle and hence will eventually dominate
the population.

Describing the optimum

There is a mixed strategy solution if
aL=kL/k<h.
Mixed solution has L =aL/h and
SL/SS= aL(1-h)/(1- aL)h.
If aL>h, then the only solution is the pure
strategy L.

Some lessons
If long winters are rare enough, the most
successful strategy is a mixed strategy.
Probability matching. Probability of Strategy L is
Is aL /h , proportional to probability of long winter.
For populations with different distributions of
winter length, but same feeding costs the die-off in
a harsh winter is inversely proportional to their
frequency.

Generalizations
Model extends naturally to the case of
many possible lengths of winter.
Replace deterministic cycle by assumption
of iid stochastic process where probability
of winter of length t is at
Choose probabilities t of storing enough
for t days. Let St() be expected survival
rate of type if winter is of length t.

Optimization
Then the optimal mixed strategy will be the
one that maximizes the product
S1() a1S2() a2 SN() aN.
Standard result of branching theory.
Application of law of large numbers. See
Robson, JET.

Do Genes Really Randomize?

Biologists discuss examples of phenotypic
diversity despite common genetic heritage.
Period of dormancy in seed plantsLevins
Spadefoot toad tadpoles, carnivores vs
vegans.
Big variance in size of hoards collected by
pikas, golden hamsters, red squirrels, and
lab ratsVander Wall

Is Gambling Better Than Sex?

Well, yes, this model says so.
Alternative method of producing variation
sexual diploid population, with recessive gene for
Strategy S.
Whats wrong with this? Strategy proportions
would vary with length of winter.
But gambling genes would beat these genes by
maintaining correct proportions always.

Casino Gambling
Humans are able to run redistributional
lotteries. What does this do?
This possibility separates diversification of
outcomes from diversification of
production strategies.
If some activities have independent risks,
individuals can choose those that maximize
expected risks, but then gamble.

A Squirrel Casino
Suppose squirrels can gamble nuts that they have
collected in fair lotteries.
Let v(y) be probability that a squirrel who collects
y days supply of nuts is not eaten by predators.
Expected nuts collected is yv(y).
Optimal strategy for gene is to have its squirrels to
harvest y* where y* maximizes yv(y) and then
gamble.

Human Gamblers
Humans are able to run redistributional
lotteries. What does this do?
This possibility separates diversification of
outcomes from diversification of
production strategies.
If some activities have independent risks,
individuals can choose those that maximize
expected risks, but then gamble.