MATH35032: Mathematical Biology, Problem Set 1
MATH35032: Mathematical Biology, Problem Set 1
(3.1)
N0 ert
(1 (N0 /K)(1 ert ))
(4) Mice in the wild obey the logistic population law (3.1). Plagues arise in mouse
populations whenever N(t) becomes too large and an epidemic can destroy 97-98%
of the population within just a few weeks. After such a disaster the mice, now
diminished to perhaps 2% of their pre-epidemic numbers, become too isolated to
permit the disease to spread. The population therefore begins to grow again until
it reaches a level susceptible to another plague.
One can model this situation as follows: say that an epidemic starts when N(t)
reaches some level Q with 0 Q < K. One then models the plague by drastically
(a) Explore the role of the parameter z by drawing curves of the form
z
N
f (N) = 1
K
where, on each curve, z is held constant and 0 N 1.1 K. Draw a
separate curve for each value of: z {0.1, 1, 10}.
(b) Determine the steady-state solutions (those with N(t) = a constant) of (5.1).
(c) Show that the equation governing small perturbations n(t) near the positive
steady-state population obey the (linearized) equation
dn(t)
n(t) (qz 1)n(t T ).
dt
(5.2)
(d) Seek solutions to (5.2) of the form n(t) = n0 exp(t) and show that any such
solution must have
= (qz 1)eT .
(5.3)
Considering C and defining = + i, find the real and imaginary parts
of (5.3).
(e) Find conditions under which it is possible that there is an exponentially growing solution to (5.3). That is, find conditions on the parameters that permit
solutions with > 0. In particular, by solving for the limiting case = 0,
show that if (qz 1) > 1, there is a critical delay Tc such that if T < Tc ,
then < 0. Show also that if (qz 1) < 1, then < 0 for all values of the
delay T .
(f) Now concentrate on the potentially unstable regime, where (qz 1) > 1, and
define
b = (qz 1).
Consider delays just slightly greater than Tc . That is, put T = Tc + , where
0 < 1, into (5.3) and, by finding , show that the period of the unstable
oscillation is
2
2
=
+ O().
b2 1
Symmetry and Snail Shells
This last problem was inspired by a set of three papers reprinted in Taubess book.
The first, (Gould 1995), is by the great naturalist Stephen J. Gould and points
out that the vast majority of know species of snails have shells that are coiled in a
right-handed sense, where right-handed is defined as described in Figure 1 below.
The other two articles (Mochizuki et al. 1998, Vogan & Tabin 1999) have to
do with the biological basis of this asymmetry: the biological reality is a lot more
complicated and subtle than models such as the one sketched below might suggest.
L
,
L+R
where L and R are, respectively, the numbers of left- and right-coiling snails.
The snails mate at a rate , so that the probability that a given snail mates
in a brief interval of length t is t.
On average, each mating gives rise to m new snails.
If two left-coiling snails mate, they produce left-coiling offspring. Similarly, a
mating between two right-coiling snails produces right-coiling offspring. But
a mating involving one snail of each type produces offspring of each type with
equal probability.
The resulting model is
d
= (1 )( 1/2)
dt
where is the fraction of left-coiling snails
(6.1)
(a) Find all the equilibria of (6.1) and classify them according to stability.
(b) Derive (6.1) from the assumptions above.
References
Gould, S. J. (1995), Left snails and right minds, Natural History 104(4), 1018.
Mochizuki, T., Saijoh, Y., Tsuchiya, K., Shirayoshi, Y., Takai, S., Taya, C.,
Yonekawa, H., Yamada, K., Nihei, H., Nakatsuji, N., Overbeek, P., Hamada, H.
& Yokoyama, T. (1998), Cloning of inv, a gene that controls left/right asymmetry and kidney development, Nature 395(6698), 177181.
URL: http://dx.doi.org/10.1038/26006
Vogan, K. & Tabin, C. (1999), Developmental biology - a new spin on handed
asymmetry, Nature 397(6717), 295.
URL: http://dx.doi.org/10.1038/16796