iSquared-Issue1 - Revista

isquared
mathematics beyond the imaginary
Autumn 2007
Extinction
modelling
Prisoners
Dilemma
John von
Neumann
THE MATHEMATICS OF
MALARIA
2.80
isquared
Volume 1 Number 1
8
Editorial
When I set out to produce this magazine, my aim was to bring
together a collection of articles that reflected the wide range of
modern-day applications of mathematics. Many people are
unaware that maths is more than just abstract concepts,
inaccessible to all but those with a university education in the
subject. In fact, mathematics can be appreciated by everyone.
The past few decades have seen maths being used in numerous
innovative real-world situations; notably in the areas of biology
and medicine, where new insights are emerging from the use of
mathematical modelling.
In this issue, we find out how maths can be used in
conservation efforts and in the battle against malaria. These are
both worldwide issues, and give some idea of the importance of
mathematics in the world today. On a lighter note, articles on
sailing and game theory highlight the potential scope of
applications. The magazine also contains a number of regular
features, including a book review, a prize crossword and several
Japanese number puzzles.
If you have any comments on the articles printed here, or
suggestions for future issues of the magazine, please write to
us or send an email. We would like to hear from you, and will
print a selection of readers letters in the magazine. Also, please
visit our website, where you can subscribe or just find out more.
Lastly, thank you for reading. I hope you enjoy this issue of
iSquared magazine.
30
Sarah Shepherd
Cover image by Eric Issele
iSquared magazine, published quarterly. Issue 1 Number 1 (ISSN 1755-7275).
Postal address: iSquared magazine, 1 Pound Cottages, Shillinglee, Godalming, Surrey, GU8 4SZ.
Email: editor@isquaredmagazine.co.uk. Website: www.isquaredmagazine.co.uk.
2 iSquared magazine
Printed in the UK by
THE MAGAZINE PRINTING COMPANY
www.magprint.co.uk
Autumn 2007
autumn 2007
contents
FEATURES
Optimisation on the high seas
By Catherine Buchanan and David Stern

Dynamic programming is a powerful mathematical tool which can be applied to
find the quickest route between two points when travelling by sailing boat.
Cover Story: Can mathematics solve

the problem of malaria?
16
By Deborah Cromer
How mathematical models can shed light on one of the worlds deadliest diseases.
The mathematics of being nice
24
By Graeme Taylor
Mutual co-operation gives the best outcome, but what is the winning strategy in
game theorys Prisoners Dilemma?
Survival or extinction?
30
By Daniel Rowe
Population viability analysis: determining whether a species will stick around.
REGULARS
News
Mathematical greats
14
The life and work of the Hungarian-born US mathematician, John von Neumann.
Book review
22
Symmetry and the Monster, the story of one of the greatest quests of mathematics.
Subscription form
29
Puzzles
36
Endnotes
39
www.isquaredmagazine.co.uk
iSquared magazine 3
news
Rubiks number
The Rubiks cube has been testing
minds around the world since its
invention in 1974. But now
scientists have come one step
closer to finding the optimal
solution to the puzzle. Daniel
Kunkle and Gene Cooperman
from Northeastern University in
Boston, Massachusetts have
proved that the cube can be solved
in no more than 26 moves.
The computer scientists came up
with the proof using brute force,
programming a supercomputer to
test various combinations. To
reduce the required computer time
to a maneagable level, they used a
two step method, where initially
the computer was programmed to
come up with 15,000 half-solved
solutions, which could be then be
fully solved easily with just a few
extra moves. The outcome of this
work showed that a cube with any
initial configuration could be
solved with a maximum of 29
moves, but that most required
only 26 moves. Kunkle and
Cooperman then used the
supercomputer to tackle those
problem cases that needed more
than 26 moves. Despite the small
number of such cases, the
calculation took the
supercomputer 63 hours. However,
the program was successful the
computer was able to solve all the
special case cubes in fewer than
26 moves.
Kunkle and Cooperman intend
to continue their research,
confident that they can further
reduce the number of moves from
26. The minimum number of
moves needed to solve any
Rubiks cube is known as Gods
4 iSquared magazine
Any Rubiks cube can be solved in a maximum of 26 moves

number named because only
God would be able to solve the
cube in so few moves. Past
research has suggested that Gods
number is in the low 20s.
simplified the model and

succeeded in developing practical
methods for solving these
equations.
The model will have important
applications for shipping safety
Sand wave dynamics are

a key factor in determining
A Dutch researcher, Joris Van Den the shape of the sea floor
Berg, has developed a
in the southern North Sea
Making waves
mathematical model for sand

waves. These are formed when the
tidal current interacts with loose
sand on the sea bed, creating wave
patterns, which themselves affect
the tidal flow and cause further
build-up of sand. The equations
describing this behaviour were
already known, but Van Den Berg
and the design of offshore

infrastructures, especially since
sand wave dynamics are a key
factor in determining the shape of
the sea floor in the southern North
Sea. It has already been used to
determine how sand waves will
recover after a trench is dredged
Autumn 2007
Mbius puzzle
A 75-year old problem involving
the Mbius strip has finally been
solved, by two researchers at
University College, London. The
Mbius strip, which was first
discovered in 1858, can be created
by taking a strip of paper, twisting
one end through 180, and then
joining the ends. The resulting
one-sided shape has fascinated
mathematicians and artists alike,
including M.C. Escher, who
famously depicted the strip in an
artwork.
Now Gert van der Heijden and
Eugene Starostin have succeeded
in defining the shape

mathematically. They were able to
use a set of 20-year-old
differential equations to describe
the very special shape of the
Mbius strip. Their research could
have applications in fabric design,
helping to predict points of tearing,
or in structural modelling during
the development of new drugs.
Tumour control
Scientists at the University of
California have used mathematics
to show why cancer cells alter
their own genetic makeup in the
early stages of tumour growth.
Natalia Komarova, Alexander
Sadovsky and Frederic Wan used
a mathematical technique called
optimal control theory to explain
biological evidence of tumour

growth patterns, observed in
laboratory studies.
Tumour cells seem to adopt a
strategy of genetic instability
Instability can have

both a positive and a
negative effect on the
rate of cancer growth
during initial development. While
this increases the probability of
cancerous mutations, it can also
cause a large death rate among the
dividing cells. Thus instability can
have both a positive and a
negative effect on the rate of
cancer growth. Komarova and her
colleagues wanted to find out
what would be the optimal growth
strategy for the tumour.
The Mbius strip has been described mathematically using differential equations
iSquared magazine 5
David Benbennick
for the planned high-voltage cable

between England and the
Netherlands.
Optimal control theory is a

method of minimising a
mathematical cost function
subject to a set of constraints on
the variables involved. The
strategy corresponding to the
minimal (or optimal) solution is
known as the control law. The
scientists in California examined
the effect of different parameters
and conditions on tumour growth.
Komarova explains: the mutation
rate serves as the control knob.
Then, we can calculate
mathematically how long it takes
a tumour with given parameters to
reach a certain size. We found
that at early stages of tumour
growth, instability is
advantageous, and later on it
becomes an impediment. This
explains why many tumours
exhibit a high level of instability
at first, and become stable later in
their development.
This research sheds light on
how cancerous tumours are able
New research has shed light on tumour growth strategies
to thrive despite a high mutation
rate, knowledge which may prove
to India in the fifteenth century.
important for cancer treatments in century, was in fact identified by
the future.
a small group of scholars in
The research team at Manchester
has even suggested that
southwest India, known as the
knowledge of the infinite series
Kerala School, in 1350. The
was eventually passed on to
beginnings of modern maths is
Indian discovery
Newton. However, Dr Joseph
usually seen as a European
Researchers at the University of
achievement but the discoveries in says: The brilliance of Newton's
Manchester have uncovered
work at the end of the seventeenth
evidence that the infinite series, Discoveries made in
century stands undiminished
one of the basic components of
medieval India have been especially when it came to the
calculus, was in fact discovered
algorithms of calculus. But other
ignored or forgotten
by Indian mathematicians 250
names from the Kerala School,
years before Newton. Dr George
notably Madhava and Nilakantha,
medieval India between the
Gheverghese Joseph has found
fourteenth and sixteenth centuries should stand shoulder to shoulder
papers showing that the infinite
with him as they discovered
have been ignored or forgotten.
series, which had previously been
There is also strong evidence
the other great component of
attributed to Newton and Leibnitz that knowledge was passed on to
calculus infinite series.
at the end of the seventeenth
Jesuit missionaries who travelled
6 iSquared magazine
Autumn 2007
Image courtesy of Dr Timothy Triche/National Cancer Institute
news
resources is a prominent problem

in environmental engineering,
Two MIT researchers have come
says Stocker. Awareness of the
up with an explanation for an
fundamental mechanisms
experiment that has baffled
governing the interaction between
scientists for many years. When
the two phases is critical to devise
an oil drop mixed with a small
sound engineering solutions for
amount of surfactant (a material
remediation. However, there are
that reduces surface tension, such also links with biological research,
as detergent) is placed on a water since spontaneous oscillations are
surface, it appears to throb like a
observed in many natural systems,
beating heart. Now Roman Stoker, for example in nerve cells and
an engineer, and John Bush, a
muscle tissue. Some scientists are
mathematician, have discovered
interested in using the oil drop
the mechanism behind this
problem to help explain such
phenomenon. Variations in
biological oscillations.
surface tension due to the
evaporation of the surfactant
cause the oil drop to alternately
Ramanujan play
expand and contract. If the
experiment is covered the
A new play about the Indian
throbbing stops, because the
mathematical genius Srinivasa
surfactant is unable to evaporate.
Ramanujan is opening at the
This work could have
Barbican Centre in London this
applications in a range of fields,
September. A Disappearing
including engineering and biology. Number, performed by
Oil contamination of water
experimental theatre company
Oil phenomenon
Joris Jan-Bos
A Disappearing Number
Complicite, explores the

relationship between Ramanujan
and the English mathematician
G.H. Hardy. The play is formed
of interlocking stories, with the
story of Ramanujan and Hardy
weaved into that of a present day
mathematician who travels to
India in Ramanujans footsteps,
and her lover, who follows after
her death.
The importance of Ramanujans
work was first recognised by
Hardy, who persuaded Ramanujan
to travel to England. In 1913
Ramanujan joined Hardy at
Cambridge, where he was plagued
by ill-health, which was
exacerbated by dietary problems
(as an orthodox Bramin,
Ramanajan was a strict vegetarian
and the First World War made
obtaining special items of food
extremely difficult). He died in
1920 aged just 33, but during his
short life made substantial
contributions in the areas of
mathematical analysis and
number theory.
Through the story of
Ramanujan and his search for
mathematical truths, A
Disappearing Number seeks to
explore the themes of
mathematics and beauty, the
nature of infinity, permanence
and continuity, love and loss,
how we relate to the past and the
future, and above all our
relentless compulsion to
understand.
A Disappearing Number is
on at the Barbican Theatre
from 5th September to 6th
October.
Sarah Shepherd
iSquared magazine 7
8 iSquared magazine
Autumn 2007
OPTIMISATION
on the high seas
Dynamic programming is a powerful mathematical
tool which can be applied to find the quickest route
between two points when travelling by sailing boat
Image by Manfred E. Fritsche (licensed by Creative Commons Attribution ShareAlike 2.0 Germany License)
By Catherine Buchanan and David Stern
he omnipresence of mathematics in our

world is reflected in this article by
considering sailing and looking at just a
few of the mathematical elements associated with it.
We look at the optimisation problem of finding the
quickest route. Optimisation is the branch of
mathematics used to find the best feasible solutions
to a huge range of problems, including organising
timetables, stocking shops and laying telephone lines.
Many optimisation tools exist, several of which have
been applied to the sailing problem. Here dynamic
programming will be used, since it is incredibly
adaptable, yet based on an easily understandable
premise that demonstrates the beauty of mathematics.
Dynamic programming was invented by Richard
Bellman in 1957. It is used to solve optimisation
problems which can be divided into stages, where
the decisions made at each stage are independent of
previous decisions. This is best illustrated by the

simple problem of finding the shortest route through
a road network with one way streets, such as that in
the diagram below.
2
3
H
2
Here, the distances between intersections are marked

on the arrows. Suppose we are required to find the
shortest route between intersection A and
iSquared magazine 9
intersection H. First, f(X) is defined to be the

shortest path from intersection A to intersection X.
Therefore
f(A) = 0,
f(B) = 5,
This method of finding the best path in a step by step

manner is more efficient than finding the lengths of
all possible paths and then choosing the shortest.
Building the sailing model
and
f(C) = f(B) + 9 = 5 + 9 = 14.
To formulate a real-life problem mathematically a

model must be built. In the simplest case, the sailing
It is possible to reach intersection D from two
model consists of a model of a boat and a model of
directions (either from A or from C), so the direct
the environment that the boat is sailing in.
route between A and D is compared with the route
A model of a boat has many different
via C, and the length of the shortest route is assigned characteristics. One essential characteristic of any
to f(D). The direction choice which leads to this
sailing model is how a boat interacts with the wind.
route is remembered. That is,
Every type of boat has its own distinct pattern of
speeds relative to the direction sailed in and the
direction of the wind.
f(D) = min{f(A) + 8, f(C) + 5}
= min{0 + 8, 14 + 5}
=8
Next the shortest route to intersection E is calculated
by considering all paths leading directly into E. This
process continues until intersection H is reached.
The details are as follows:
This can be seen most easily in the form of a wind
polar. Wind polars are polar coordinate plots of a
boat's relative speed (r) versus the angle made with
the wind ().
f(E) = f(D) + 3 = 8 + 3 = 11,

f(F) = min{f(C) + 3, f(E) + 2}
= min{14 + 3, 11 + 2}
= 13,
f(G) = f(F) + 3 = 13 + 3 = 16,
f(H) = min{f(E) + 8, f(G) + 2}
= min{11 + 8, 16 + 2}
= 18.
These solutions can be seen in the following diagram,
where the shortest routes from A to each intersection
are shown. The best route from A to H is via D, E, F
and G, and is 18 units long.
14
10 iSquared magazine
11
13
18
16
Two examples of
wind polars are
illustrated in the
diagrams above
and right. The top
wind polar is
taken from Robert
Vanderbei's
intuitive guess at
boat speeds with
respect to the
wind. However,
intuition can often be misleading and there is no
substitute for accurate data. The wind polar on the
Autumn 2007
right was found by practical experimentation. Mike

Hennessey and colleagues carried out experiments
with a C&C yacht on Lake Superior. They used a
wind sensor, GPS data logging and specially
designed computer software in order to gain
sufficient data to determine the wind polar.
Neither of these wind polars corresponds to what
the authors experience of sailing led us to expect.
We expected a boat to travel at similar speeds when
travelling with between 0 and 90, unlike the first
polar. Looking at the second polar we were
surprised at how much more slowly a boat travels
with the wind than when travelling at a small angle
with it.
More experienced sailors have confirmed that the
second wind polar is actually accurate. This implies
that tacking is useful when travelling downwind as
well as when travelling upwind. Tacking is the
process of turning the sail across the wind, and
invariably causes a loss of speed which can be
modelled by associating a tacking penalty to the boat
model.
Essential components of the environment model
include a region in which the boat can travel and
wind data, which is comprised of both direction and
strength. Obtaining accurate data to apply this model
to a real-life problem is already complex. For
example, to define a precise model for the region in
which a boat sails, one would require detailed
survey maps of the water, and to formulate a model
of the wind complicated meteorological data for
everywhere in the region would be needed.
In this article we will associate only these two
components to the model of the environment,
assuming simplistic wind data and approximating
water regions with simple shapes. However, the
methods presented here are not limited to this model.
It can be extended as far as your imagination will
take you.
Dynamic programming for the

sailing problem
In practice, the direction in which a boat is sailed
can be continuously adjusted. However, dynamic
programming is a technique which is applicable to
discrete problems, so the sailing route is divided in
such a way that decisions are made at discrete time
intervals of 1 unit. Also, the choice of sailing angle
with the wind () is restricted, so that is allowed to
take a limited number of specified values between

-180 and 180.
The function f(s,x1,x2,w) is defined to be the
minimum time taken to reach position (x1,x2) at stage
s, with the sails set to the w side of the wind (R =
right or L = left). Keeping a record of the side of the
wind to which the sails are set enables us to impose
a tacking penalty every time a route crosses over the
wind.
We will illustrate the use of dynamic programming
with a simple problem that can be worked through in
detail. It consists of sailing from the west to the east
side of a lake with a constant south westerly wind of
strength 10. The lake is approximated by a square,
with sides of length 20 units. It is represented by a
coordinate system (x1,x2), with the western bank at
x1 = 0 and the southern bank at x2 = 0.
First we must define a wind polar for our boat. For
the purpose of demonstration, only 5 sailing
directions are allowed:
The tacking penalty chosen is to halve the boat's

speed during a stage when it tacks.
STAGE 0
Our boat is placed at the centre of the western shore
of the lake. Two representations of the boat are
considered; one which has its sails set ready to sail
to the left of the wind and one which is ready to sail
to the right of the wind. The starting points are as
follows:
f(0,0,10,R) = 0 and f(0,0,10,L) = 0.
STAGE 1
From the starting position, 4 of the possible 5 sailing
directions keep the boat on the lake. The positions
reached by sailing in these directions are illustrated
on the following page.
iSquared magazine 11
them. Part of the challenge of creating a dynamic

programming algorithm is to find a way of
eliminating positions without compromising the
accuracy of the model. A simple suggestion is to
take each possible position in turn and eliminate any
which are within a given radius of a position that has
already been considered. However, there are many
other ways of reducing the number of possible
positions and comparing solutions obtained from
such different algorithms can be as interesting as
comparing different models.
Sailing models using a computer

The figure clearly shows that two positions are
reached in each of the four sailing directions. This is
because the boat only travels half as far during a
stage when it tacks.
STAGE 2
From the 8 positions reached at stage 1, another 31
positions can be reached (13 of these can be reached
in two different ways). It is the recognition that each
time a position is reached in more than one way it
can be considered as just one stage that is the key to
the power of the dynamic programming technique.
The positions reached are shown below.
Even after eliminating some positions the problem

size increases so rapidly that it is almost
immediately necessary to solve it using a computer
rather than by hand.
Recall the two wind polars considered earlier, for
Vanderbeis imaginary boat, and the C&C yacht.
Our modelling gives us a chance to compare these
boats in different winds. We assume that the top
speeds of these boats are equal but obtained when
sailing at different angles to the wind.
Sailing with the wind:
The first problem is to find the fastest route for
sailing a boat from a starting point at x1 = 0, x2 = 35
(S) to a finish line at x1 = 400 (F) along a river
which is 70 units wide. There is a constant following
wind (from the west) of strength 35. Typical optimal
routes for each boat are shown in the diagram below.
The eastern bank of the lake is reached at this stage

with f(2,20,10,R) = 2. Therefore, the calculations
The imaginary boat, whose fastest speed is attained
can stop here with the conclusion that the optimal
when sailing with the wind, would sail down the
route for this sailing boat to cross the lake takes 2
centre of the river for 12 units of time, whilst the
units of time and is achieved by sailing at an angle
C&C yacht would tack from one side of the river to
of -45 to the wind for the duration of the journey.
The number of possible positions increases rapidly the other, taking 17 units of time to reach the finish
line.
at each stage and it is not useful to work with all of
Autumn 2007
Sailing in a sinusoidal wind:

In our second problem the direction of the wind
varies continuously with x1. This is illustrated by
plotting the angle of the wind against the distance
traveled across the lake:
The wind has a constant strength of 35. An optimal

route from a starting point at x1 = 0, x2 =100 (S) to a
finish line at x1 = 400 (F) along a river which is 200
units wide is sought. Typical optimal routes for each
boat are shown below. Here, the changes in the
angle of the wind as the boat crosses the lake are
represented by arrows above the diagram.
The boats would take similar routes, travelling in a

south easterly direction for the first half of the
journey, when the wind is from the north, then
tacking and continuing in a north easterly direction
for the second half of the journey when the winds
are from the south. In a C&C yacht, the journey
would take 28 units of time whilst in the imaginary
boat it would take 44 units of time.
Understanding or teaching sailing can be helped by
these very simple models which demonstrate the
effects of specific properties of the boat or the
environment. In the sailing with the wind example
given above, the difference in wind polar has a large
effect on the route that the boat should take. In the
second problem it is interesting to observe that
although the two wind polars are very different, they
react to changing wind in a similar way.
At every stage of the modelling process extra
detail could be added to edge towards a real-world
model. Each addition in itself is not fundamentally
complicated, but could require considerable time
and/or knowledge to implement. An improved realworld model could be useful for boat design, in
ensuring safety at sea or even to create that ultimate
pirate game that you have always wanted.
About the Authors
Cathy Buchanan studied mathematics at
Sheffield University, before working in a variety
of jobs, including stock control, teaching English
abroad and planning delivery schedules. She is
now a postgraduate student at the University of
Edinburgh, where her research area is
operational research.
David Stern grew up in West Africa, but
moved back to the UK to work as a programmer.
Later he took an undergraduate mathematics
degree at Warwick University, and went on to
study pure mathematics at postgraduate level,
first at Edinburgh University, then Sheffield. His
current research is related to string theory.
FURTHER READING
Dynamic Programming. Richard Bellman, Princeton University Press, 1957.
Integrated graphical game and simulation-type problem-based learning in kinematics. M. P. Hennessey and S.
Kumar in International Journal of Mechanical Engineering Education, Vol. 34, No. 3, pages 220-232, 2006.
Optimal sailing strategies, statistics and operations research program. Robert J. Vanderbei, University of Princeton, http://www.sor.princeton.edu/~rvdb/sail/sail.html, 1996.
Optimal routing of a sailboat in steady winds (unpublished). M.P. Hennessey, J.A. Jalkio, C.S. Greene and C.M.
Sullivan, School of Engineering and Centre for Applied Mathematics, University of St. Thomas, 2006.
mathematical greats
The life and work of the Hungarian-born US mathematician, John von Neumann
Born in Budapest in 1903, John

von Neumann was the eldest child
of a Jewish banker. He was
quickly recognised as a child
prodigy. Aged 6, he was able to
divide two eight-digit numbers in
his head, and by 8 he had
mastered calculus. As a child he
would sometimes amuse family
guests by memorising the pages
of phone books on sight. In 1911
he entered the Lutheran
Gymnasium in Budapest, where
his genius was soon recognised.
Von Neumann completed his
schooling in 1921 and the
following year published his first
mathematical paper. However, his
father wanted him to study a
subject with better financial
prospects, so as a compromise he
agreed to study chemistry at
university. Having also won a
place at the University of
Budapest to study mathematics,
he received both a PhD in
mathematics and a diploma in
chemistry (from ETH Zurich in
theories was one of the 23 famous
Switzerland) at the age of 23.
problems set by David Hilbert at
For the next four years von
the International Congress of
Neumann taught at the University Mathematics in 1900, and while at
of Berlin, during which time his
Berlin von Neumann started
fame spread rapidly in the
working on building a solid
mathematical community. His
mathematical framework for
doctoral thesis had been on set
quantum mechanics, whichat that
theory, and he continued to work
time was still a very new theory.
in pure mathematics, in the areas
In the 1930s, after Hitlers rise
of logic and analysis. He was
to power, von Neumann
particularly concerned with
emigrated to the US with his
building a solid foundation for
mother and brothers. He had been
mathematics, by defining the
offered a job as mathematics
basic axioms from which all
professor at the Institute of
theorems can be derived. The
Advanced Study in Princeton. He
axiomatisation of physical
was one of the first six members
Image courtesy of Los Alamos National Laboratory
If people do not believe that mathematics is simple, it is only because

they do not realise how complicated life is. John von Neumann
of the newly formed department

(along with Einstein), and worked
there from its formation until his
death. At Princeton, he continued
to work in quantum mechanics,
publishing a book in 1932 in
which he laid out mathematical
foundations for the physical
theory, using the concept of
rings of operators, now known as
Neumann algebras.
Von Neumann was in many
ways the antithesis of the
stereotypical mathematician. He
was hedonistic, tended to dress
formally and loved throwing wild
parties. He was also a very bad
Autumn 2007
driver, and reportedly often read a end of the Second World War his
book while driving. An account he support for the development of
gave of one car accident was: I
nuclear weapons remained
was proceeding down the road.
unperturbed. He once described
The trees on the right were passing his own political ideology as
me in orderly fashion at 60 miles violently anti-communist, and
per hour. Suddenly one of them
much more militaristic than the
stepped in my path. In 1930 von
norm. This has led to some
Neumann married his first wife,
negative opinions of him. Indeed,
with whom he had one child, a
it has sometimes been said that
daughter named Marina. It is said von Neumann, who was confined
that when he made his proposal of to a wheelchair in later life, was
marriage, all he could come up
the model for the delusional title
with was: You and I might be
character of Stanley Kubricks
able to have some fun together,
1963 film Dr Strangelove.
seeing as how we both like to
However, von Neumann also
drink. They divorced in 1937 and worked on many other areas of
von Neumann married again the
applied mathematics. He is
following year.
perhaps best known today as
With the outbreak of World War being the founder of game theory,
II in 1939, von Neumann, like
which has been widely applied to
many other mathematicians at that
time, became interested in applied
Von Neumann was a
mathematics. He developed an
hedonist who loved
expertise in explosives and was
taken on as a member of the
throwing wild parties
Manhattan Project the WWII
project to develop the first nuclear economics. When his book,
weapon. He played an important
Theory of Games and Economic
role in designing the explosive
Behaviour, which he co-authored
lenses needed for the atomic bomb. with Oskar Morgenstern, was
These compressed the plutonium
published in 1944, it received so
core evenly by focusing the
much public attention that the
diverging shock waves into
New York Times did a front page
converging waves, leading to a
story.
highly efficient nuclear explosion.
The other area of research where
Von Neumann also discovered that von Neumann made several
large bombs are more devastating lasting contributions was
when detonated above the ground computer science. The Monte
due to the force of the shock
Carlo method, which allowed
waves. In 1945 when the first
complicated problems to be
atomic weapons were dropped on approximated using random
Hiroshima and Nagasaki, they
numbers, was partly developed by
were detonated at the very altitude von Neumann. He was also
that von Neumann had calculated
involved in the conception of
would do the most damage.
single-memory computer
Von Neumann held very rightarchitecture, which is now
wing political views, and after the commonly known as von
Neumann architecture and is used

in almost all present day
computers.
Von Neumann had a strong
belief in mathematical methods
and models, and attempted to
develop a mathematical theory of
the life sciences. While working
on self-replicating systems, he
came up with the first cellular
automata. These are discrete
mathematical models consisting
of a regular grid of cells, with
each cell in one of a finite number
of states. At each time step, each
cell will change state according to
a rule dictated by the states of the
cells in the immediate neighbourhood. Neumann proved that a
particular pattern of states would
endlessly replicate itself within
the cellular universe.
In 1957 von Neumann was
diagnosed with cancer, possibly
caused by exposure to
radioactivity during his work on
nuclear weapons. He was
devastated by his illness, which
he realised was incurable. The
cancer spread to his brain and
affected his cognitive abilities.
His friend Edward Teller said of
his final days, I think that von
Neumann suffered more when his
mind would no longer function,
than I have ever seen any human
being suffer. He died within a
few months of the initial
diagnosis.
By the time of his death, von
Neumann had published 150
papers and made a lasting
contribution to a broad range of
mathematical disciplines. He will
undoubtedly be remembered as
one of the greatest and most
influential mathematicians of the
20th century.
Sarah Shepherd
Can mathematics
solve the problem of
MALARIA?
A
long with HIV/AIDS and tuberculosis,

malaria is one of the World Heath
Organisations Big 3, the three most
deadly diseases worldwide. Malaria is responsible
for 1.3 million deaths per year, and in fact only 10
years ago still caused more deaths than HIV/AIDS.
Around half of the worlds population is exposed to
malaria and between 300660 million malaria
episodes occur annually, almost all in Africa, South
America and South East Asia. The majority of those
who die from these infections (nearly 85%) are
children under the age of 5 in some of the poorest
countries in Africa. Although the disease affects
nearly half the worlds population, surprisingly little
is known about its dynamics. Mathematical models
of malaria can help to understand the disease
dynamics and thus aid in the eradication of the
disease. In this article we will take a look at the
disease itself and how mathematics can help us to
better understand it.
Malaria is a parasite which lives and grows inside
red blood cells. The parasite is injected into a patient
through a mosquito bite from an infected mosquito
(the vector for the disease). After a brief period of
By Deborah Cromer
multiplication inside liver cells, thousands of
parasites are released into the blood stream. These
invade red blood cells and there grow from a small
single parasite into large schizont, which contains
8-32 new parasites. After 48 hours the infected red
blood cell ruptures and releases the new parasites
into the blood stream, allowing the process to be
repeated.
As you can imagine, this has a devastating effect
on the patient. When the infected cells rupture they
tend to do so all at the same time, which means that
a large number of parasites are released into the
blood stream all at once. In an attempt to fight and
kill them, the patients immune system develops the
characteristic fever associated with malaria. As the
parasites will rupture every two days, the fever too
will break out every other day.
Parasitised cells prefer to live outside of the
regular blood stream, as this makes them harder to
detect. They often reside in tissues within the brain,
where they can cause coma and even death. This is
known as cerebral malaria. Another symptom
associated with the disease is severe malarial
anaemia, which is due to the fact that the parasites
Autumn 2007
Images courtesy of CDC (Centers for Disease Control and Prevention)
How mathematical models

can shed light on one of the
worlds deadliest diseases
Top: a female Anopheles mosquito feeds on a human host

Bottom: scanning electron micrograph (SEM) of an Anopheles mosquito
grow inside red blood cells, inevitably leading to the

death of the infected cells when they rupture.
Blood cell models

The majority of models to date have considered how
red blood cell populations change over time, and are
thus most applicable to this last clinical effect. We
will be concerned with such models in this article.
However, models addressing other effects have also
been published, for example on the relationship
between parasite release and fever.
Mathematicians can set up models describing the
growth and death of red blood cells and parasites. A
model can be used to predict how the number of
parasitised cells in a patient changes during an
infection, and the results from the model compared
with real-life data from infected individuals. The
model could also be used to discover the effects of
different treatments, immune responses or parasite
strains. For example, suppose a parameter in the
model governs how easily a parasite can invade a red
blood cell. Then by altering the value of that
parameter it would be possible to analysis the effect
Newly created cells

replace aging cells
Unparasitised
cells
u(t)
that having a stronger parasite has on the outcome of

the infection.
The first mathematical model of malaria was
presented by Roy Anderson and colleagues at
Imperial College London in 1989. They modelled the
effect of the malaria parasite on red blood cells by
considering how three populations of cells change
over time (see diagram below). The three cell
populations are unparasitised (healthy) red blood
cells, parasitised red blood cells and the free
parasites themselves. These are denoted by u(t), p(t)
and m(t) respectively, and are all functions of time, t.
(The letter m stands for merozoites, the technical
name for the parasite).
Uninfected cells are created at a constant rate and
live for an average of 120 days. The creation of new
cells simply replaces older cells as they die off.
Unparasitised cells become parasitised by coming
into contact with a parasite, and the probability that
such contact results in the formation of a parasitised
cell is denoted by . This means that parasitised cells
are created at a rate of u(t)m(t). Parasitised cells
have an average lifetime of 2 days, as they rupture
48 hours after being created. When a parasitised cell
ruptures it creates a number of free parasites (usually
Life cycle of red blood cells in a malaria patient
Unparasitised cells become

parasitised by combining
with a free parasite
Parasitised
cells
p(t)
Rupture of parasitised
cells creates 832
new free parasites
Unparasitised cells
die after 120 days
due to ageing
Free
parasites
m(t)
Free parasites die within 20

minutes, if they do not invade
a red blood cell
Autumn 2007
this number is between 8 and 32). The free parasites

cannot live very long outside of a red blood cell and
so either go on to invade another cell or die after an
average of 20 minutes.
This model can be described mathematically by
three coupled differential equations (one for each of
the cell populations), from which mathematicians
have made a number of observations. Simulations
of the model exhibit oscillations in both the
numbers of parasitised and unparasitised cells (see
graph on next page), which dampen towards an
equilibrium level. Also, solving the model under
the assumption that all populations are at
equilibrium, shows that the malaria infection will
take hold within a patient only if certain conditions
on are met.
The majority of deaths from malaria occur

in Africa children below the age of five
Map image courtesy of CIA Factbook
Whilst this is an interesting model which can shed

some light on the necessary conditions for an
infection to take hold, the results obtained do not
always match well with data gathered from patients
with malaria. It is difficult to find values for the
parameters with which the model gives realistic
results. In patients with malaria, the number of
parasitised cells tends to grow rapidly to levels
where they can be detected in a patient, but does
not generally rise to above 5% of the total red blood
cell pool. If the parameter in the model is high
Fotolia
Beta paradox
The worldwide distribution of malaria: dark colouring indicates high risk regions
enough to generate rapid parasite

growth, far too many cells become
parasitised. However if we set
low enough so that only a realistic
number of cells become
parasitised, then it takes too long
for the number of parasitised cells
to reach a level where they would
be detectable in patients.
An obvious reason for this
paradox is that there has not yet
been a term included in the model
to account for any immune
response which a patient mounts
against the infection. It is almost
certain that in an attempt to
combat the disease there is some
killing of infected cells by the
patients immune system, although
the exact form that this takes is as Results from a simulation of the basic malaria model, where
yet unknown. This is an example 100 is the number of cells present in a healthy, uninfected
of how a model that does not
person. After initial oscillations, the cell populations settle
exactly fit the data can give some down to steady levels.
insight into an important aspect of
the disease. From the results
1) How are the T-cells created? Is the number
obtained using the model, we can infer that the
of T-cells created dependent on the number
immune response during a malaria infection is
of parasites already in the system? Or on free
important, since omitting it means that the model
parasites? Or perhaps on both?
cannot accurately fit the data.
2)
Does
the number of T-cells created depend
One of the advantages of creating a mathematical
on the number already present (i.e. can they
model of biological phenomena is that it is possible
help to generate themselves)?
to hypothesise about various aspects of the system
3) How do the T-cells kill the parasitised cells
that we think might be important and include these in
and free parasites?
the model. We can then decide whether the results
Answers
to each of these questions result in new
obtained from the new model better match with the
experimental data. In the case of a malaria infection, assumptions being made about the model, and also
lead to the inclusion of new parameters.
this involves including an immune response which
After the modellers made the decisions above, it
takes on a biologically reasonable form.
turned out that an additional six unknown parameters
needed to be included in the model. Although this
new model is more realistic, and can be used for
Improving the model
more complex inferences, there are also many more
An issue faced when adding any new aspect to a
unknowns. This means that there are a wide variety
model is that of the corresponding new parameters,
of different model solutions, depending on the values
which add to those already present and serve to
of the six extra parameters.
complicate the mathematics. Modellers have added
Despite this caveat, by extending the model to
an immune response to the basic model described so include an immune response the modellers were able
far. The immune response is effected by a fourth cell to conclude that immunity targeted against the
population, T-cells. These T-cells perform the killing parasitised cells would be more effective than an
of parasitised cells and free parasites. Before adding immune response which killed off free parasites.
the T-cells into the model a number of decisions had This is an important observation for researchers
to be made:
developing vaccines against malaria.
Autumn 2007
In general there is a balance to be achieved

between adding complexity to a model, and therefore
making it more biologically accurate, and reducing a
model so that it contains fewer unknown parameters.
Each scenario is different, and depends on both the
system being modelled and the data available from
which to estimate parameters.
Many other extensions of the basic malaria model
have also been published. In 1992, Barbara
Mathematical modelling provides

a safe and inexpensive way to
explore how the infection behaves
fitted a slightly different model of a malaria infection

to patient data. Using their model they were able to
provide further evidence to biologists that this extra
killing does occur, and estimated that on average 8.5
healthy cells are destroyed for every one cell that
becomes parasitised. Their work implies that one of
the major causes of severe malarial anaemia is the
extra killing of healthy cells, and this insight can
influence the treatment of patients with the disease.
Mathematical modelling of malaria is just one of the
tools that can be used in the fight to understand and
ultimately eradicate this disease. Modelling provides
a safe and inexpensive way to explore various
aspects of how the infection behaves. In this article
we have explored the complex interactions of red
blood cell production, destruction and parasitisation
which take place in patients with malaria. Different
models are also being developed to address other
aspects of infection. With the help of the improved
understanding that mathematics is able to shed on
malaria we can hope that one day the WHOs Big 3
will be reduced to the Big 2.
Hellriegel, a zoologist at the University of Basel,

presented an extended model which considered the
interaction between two different strains of the
malarial parasite. This is especially relevant for
Africa, where there are many different forms of
malaria of varying severity, and patients are often
infected by new strains whilst still fighting an old
infection. The interactions between species are by no
means simple and modelling suggests that the long
term outcome depends largely on which strain
About the Author
infected the patient first and when the second one
Deborah Cromer is a
appeared. In fact, it turns out that infection with a
PhD student at Imperial
mild strain of malaria early on can have a
vaccinating effect, actually reducing the symptoms if College. She completed
her undergraduate
a more virulent strain infects a patient later.
degree at the University
Another variation of the model was developed by
of New South Wales in
G. Jakeman and other researchers in Queensland,
Sydney, Australia, before
Australia, who were interested in determining
going on to work for a
whether there was any extra killing off of healthy
medical research
cells during a malaria infection. This phenomenon
company, designing
has been suggested by biologists and could be
algorithms to detect
because healthy cells are mistaken for parasitised
cancer. A Mathematical Biologist, her current
cells, or because the body is consciously removing
work is on malaria, within both the human host
red blood cells so that there is nothing left for the
and the mosquito.
parasite to invade. The modellers in Queensland
FURTHER READING
Periodic and chaotic host-parasite interactions in human malaria. D. Kwiatkowski and M. Nowak in Proceedings
of the National Academy of Sciences USA, Vol. 88, No. 12, pages 5111-5113, 1991.
Non-linear phenomena in host-parasite interactions. R.M. Anderson, R.M. May and S. Gupta in Parasitology, Vol.
99, Supplement, pages S59-79, 1989.
The within-host cellular dynamics of bloodstage malaria: theoretical and experimental studies. C. Hetzel and
R.M. Anderson in Parasitology, Vol. 113 (Pt 1), pages 25-38, 1996.
Modelling the immune response to malaria with ecological concepts: short-term behaviour against long-term
equilibrium. B. Hellriegel in Proceedings of the Royal Society, Series B, Vol. 250, No. 1329, pages 249-256, 1992.
Anaemia of acute malaria infections in non-immune patients primarily results from destruction of uninfected
erythrocytes. G.N. Jakeman et al. in Parasitology, Vol. 119 (Pt 2), pages 127-133, 1999.
book review
Building blocks for the universe?

Symmetry and the Monster:
The Story of One of the
Greatest Quests of
Mathematics
by Mark Ronan
OUP 2007, 8.99 (Paperback)
Mathematicians love patterns.
Indeed, mathematics is sometimes
defined as the study of patterns.
And so it is logical that the largest
collaborative project of all time in
mathematics should be the quest
to find the fundamental building
blocks of symmetry, that most
important component of
mathematical patterns.
Mark Ronan begins this
exploration of symmetry with the
work of the ancient Greeks,
describing how the five Platonic
solids the tetrahedron, cube,
octahedron, icosahedron and
dodecahedron were discovered,
and examining their symmetries.
Leaping forward in time to the
nineteenth century, we are
introduced to the young genius
variste Galois, who famously
died in a duel at the age of just 20,
leaving behind some
groundbreaking work on the
symmetry of solutions to algebraic
equations. The objects examined
by Galois were systems of
permutations, or groups. This set
the scene for the symmetry quest:
in Galoiss work a vital
component was the idea of
deconstructing a group into
simpler groups, and it is these
atoms of symmetry, groups
which can be deconstructed no
further, that mathematicians

wanted to discover.
The next key character in the
narrative is Sophus Lie, a
Norwegian mathematician who set
out to do for differential
equations what Galois had done
for algebraic equations. The
resulting Lie groups of
continuous transformations were
to form the basic models for most
of the atoms of symmetry. By the
start of the twentieth century,
Lies groups had been classified,

and had then been telescoped
down from the continuous world
into a periodic table of finite
symmetry atoms.
However, the mathematicians
knew that there existed some
exceptional symmetry atoms,
which were not found in the
periodic table. The first
exceptions had been discovered in
the mid-nineteenth century by a
mathematical physicist named
The icosahedron an object with a very special symmetry

Autumn 2007
mile Mathieu. Much later, in

Moonshine, and although it has
1963, when mathematicians were been proved that the relationship
making detailed studies of the
is more than coincidental, the
families of symmetry atoms in the underlying reason for this
periodic table, another exception
connection remains elusive.
was found by Zvonimir Janko.
There are also deep mathematical
Jankos discovery really set the
links to particle physics it has
cat among the pigeons. No longer been demonstrated that the
could one assume that the periodic Monster is the symmetric group of
table, along with the groups of
a special string theory. This may
Mathieu, was a complete list of
indicate that the Monster is
symmetry atoms.
somehow involved in the
There then followed a spate of
fundamental structure of the
similar discoveries. New
physical world. Understanding
exceptional symmetry atoms were
found by Janko, Michio Suzuki,
It was calculated that
John Conway, Bernd Fischer and
the Monster group
others. Many of these were huge
could only exist in
in size and complexity, but the
largest of all were yet to come. In
196,884 dimensions
1973, Fischer discovered two
more groups, named the Monster
its full nature is likely to shed
and the Baby Monster. The
light on the very fabric of the
smaller of the two, the Baby
universe.
Monster, had size
Ronan brings across an
4,154781,481,226,426,191,
infectious enthusiasm for the hunt
177,580,544,000,000, yet the
for the Monster, portraying the
Monster was far greater. But did it excitement of mathematical
actually exist?
discovery and a personal love of
Mathematicians were daunted
research: Mathematics will never
by the immensity of the task of
be fully known. There will always
constructing the Monster. When
be deeper levels to uncover and
Fischer was working on the
further surprises in storeit is a
Monster he reckoned that the time subject that compels creativity,
required for just one matrix
driving mathematicians forward
multiplication would be about half on quests that are beyond the
a year of computer time. But in
power of any individual. He
the late 1970s the Monster was
blends mathematical detail with a
constructed by Robert Griess,
delightful selection of historical
an American mathematician.
facts and anecdotes about the
It was calculated that the
people involved in his narrative.
Monster group could only exist in For example, he recounts the story
196,884 dimensions.
of Lies arrest and imprisonment
Astoundingly, a connection was as a suspected German spy while
discovered between the Monster
hiking in Italy. His mathematical
and number theory, a separate
papers were taken to be coded
mathematical discipline. This has
messages, lines and spheres
been dubbed Monstrous
being interpreted as infantry
and artillery.
During the course of the book,
Ronan discusses a variety of
mathematical fields, from pure
subjects such as group theory and
number theory, to the physical
theories of relativity and quantum
mechanics. His explanations of
key mathematical concepts, such
as groups, modular (or cyclic)
arithmetic and multi-dimensional
objects are well thought out and
he avoids over-complicating the
mathematics involved, retaining
only the bare necessities for
explaining the quest for symmetry
atoms.
This is a narrative of many
strands, which at times can be
confusing. Ronan too often leaves
fragments of one story
temporarily unfinished in order to
update us on another development
in the quest. However, he just
about manages to hold it all
together; no small feat with such a
large number of characters and
intertwined discoveries to make
sense of.
Scattered through this book are
morsels of information that shed
light on the process of
mathematical research the initial
ideas and their development
(people sit and talk, perhaps with
a chalkboard at hand, and as they
talk they clarify their own ideas)
through to the publication of
research papers and even the
awarding of the most prestigious
prize for mathematical research,
the Fields medal.
In short, this is a fascinating
book that will appeal to anyone
with an appetite for exploration
and discovery, and which is
accessible to all.
Sarah Shepherd
THE MATHEMATICS
OF BEING NICE:
building trust in the Iterated
Prisoners Dilemma
Mutual co-operation gives the best outcome,

but what is the winning strategy in game
theorys Prisoners Dilemma?
By Graeme Taylor
wo prisoners are independently interrogated

by the police for a crime they are guilty of, to
which they may individually either confess
or deny. The authorities lack firm evidence to
convict either prisoner of this crime, so if neither
confesses they can only receive a minor jail
sentence for previous crime. If both prisoners
confess to the crime, then there is no doubt over
their guilt and each receives the standard sentence.
However, as an incentive to confess, the prisoners
are told that if they confess whilst the other denies,
then they will walk free for their honesty whilst the
other is made an example of, receiving a still
harsher sentence.
This is a typical formulation of the Prisoners

Dilemma, which is an example of a strategic
game studied by game theorists. As a game, the
Prisoner's dilemma is deceptively simple there
are just two players, each with a single choice, to
either cooperate or defect. But the question of
whether cooperation can be ensured (for mutual
benefit) is a fascinating one.
Autumn 2007
The possible outcomes of

the dilemma are given in
the following table:
Prisoner A
defects
In order to analyse the

game, it is useful to convert
Prisoner A
the time served in prison
co-operates
into points won in the game.
A short sentence will correspond
to a high score for that player, and vice
versa. Converting prison time into game
scores, we can represent the scenario
mathematically as a bimatrix:
(1,1) (4,0)
(0,4) (3,3)
Prisoner B
defects
Prisoner B
co-operates
Each serves 4 years
Prisoner A serves 1 year,

Prisoner B serves 5 years
Prisoner A serves 5 years,

Prisoner B serves 1 year
Each serves 2 years
Here row one denotes a confession of guilt

by Player A (defection) and row two
denotes a denial of guilt by Player A
(referred to as the strategy of co-operation,
in the sense that a prisoner co-operates
with the other rather than with the
authorities). Similarly, the first column
represents defection by Player B, and the
second co-operation by Player B. Each
pair indicates the payoff to each player: for
instance, (4,0) indicates a score of 4 for
Player A and 0 for Player B. Here, the
payoff may be interpreted as years of
freedom over the next 5 years.
We can analyse the game from the
perspective of Player A. In the absence of
knowledge of Player B's strategy, it is clear
that Player A should defect, since row 1
dominates row 2 as follows. Supposing
Player B defects, then Player A is better
served by also defecting both are found
guilty of the crime, but Player A doesn't
incur the penalty for failing to confess.
Meanwhile, should Player B have cooperated, Player A can secure his freedom
(and hence the greatest payoff) by
defecting.
The situation is exactly the same for Player
B, who therefore must also defect (this is
reflected in the payoff matrix by column 1
dominating column 2). Thus we arrive at
the scenario of mutual defection, with a
payoff to each of 1.
Yet there is a sense in which this
behaviour is irrational. Even if motivated
entirely by self-interest, the prisoners would prefer

people choosing to co-operate more than this gamethe outcome of mutual co-operation to mutual
theoretic analysis would predict. A recent
defection (a payoff of 3 each, instead of 1 each).
experiment of this kind with university students
This outcome also corresponds to the greater good,
revealed that defection rates of non-economics
since the sum of payoffs is greater for mutual comajors was under 40%. Whilst economics majors
operation (6) than for exploiting the other prisoner
defected 60% of the time in the standard game,
to secure your freedom (4). The problem is that
when given the opportunity to make (non-binding)
neither may deviate from the defection strategy
deals with the other participants before play, both
alone, for to do would get them an extra year in jail, categories dropped to a defection rate of around
whilst their partner dodges a sentence. To secure the 30%. If one seeks to use game theory to explain
benefits of co-operation, the prisoners require some
behaviour of individuals, these discrepancies
binding arrangement. Otherwise a prisoner, acting in between theory and practice must be resolved.
accordance with self-interest, maximises their
personal gain at the cost of the group: by promising
to co-operate to secure the other participant's coThe Iterated Prisoner's Dilemma
operation, then defecting anyway to collect the
A simple change of the game fundamentally alters
higher payoff.
the rational behaviour, and hence offers some clue
To describe mutual defection as the rational
as to motives that may influence choices in the
strategy is to take the non-cooperative view of
standard Prisoner's Dilemma. By playing a larger
games. In this interpretation, players are either
game, the Iterated Prisoner's Dilemma (IPD),
unable to negotiate with each other, or simply
unable to place enough trust in the others promises consisting of several rounds of the Prisoner's
Dilemma, actions in a given round will have
to risk being short-changed by a deceptive partner.
repercussions for future play and hence for your
Each is therefore best served by a strategy which
long-term score. Thus participants have an incentive
avoids the potential for exploitation, at a probable
to co-operate early to build trust and benefit from
cost of being able to co-operate for mutual gain.
mutual co-operation later, and thus to co-operate in
Such a model
any single play
of rationality in
of the Prisoner's
non-cooperative
Dilemma.
games is
In fact, a
captured by the
strategy
along
celebrated Nash
this line turns
equilibrium, a
out to offer
pair of
greater expected
strategies such
payoff than
that neither
persistent
player gains by
defection; counilaterally
operation can
deviating from
emerge from a
their role.
non-cooperative
Surprisingly,
game.
it is possible for
co-operative
In an iterated
behaviour to
game in which
emerge from a
each round
system where it
consists of a
is not a priori
play of the
assumed by the
prisoners'
players. There
dilemma against
is considerable
the same
experimental
opponent, the
evidence for
Autumn 2007
Tit-for-Tat strategy is:

In round 1, co-operate.
Inroundnforn2,playyouropponent's
strategy from round n1.
Around 1980, political scientist Robert Axelrod
organised computer simulations of the iterated
prisoner's dilemma by inviting game theorists to
supply programs implementing their strategy of
choice. Each program was run against all the others,
itself, and a random program which opted for cooperation or defection with equal probability each
round. In a preliminary event, Anatol Rapport's Titfor-Tat program only achieved second place, with
victory going to look ahead, a program which
employed tree-searching techniques similar to those
Niceness has a massive

advantage, but being too
nice leads to exploitation
used in computer chess programs. None the less it
captured the interest of many of the participants who
sought to improve upon Tit-for-Tat for the main
competition. However, it transpired that the most
elegant formulation was also the most effective, with
classic Tit-for-Tat scoring higher than any other in
numerous follow-up events.
The remarkable feature of Tit-for-Tat was that it
would, over the course of a competition, defeat the
always defect approach, even though it would
always lose to the latter in a one-on-one situation!
Always defecting is known as an evolutionary stable
strategy, meaning that in a population of such
programs, any deviation away from the approach
(by co-operating at some time) can only
disadvantage you. Tit-for-tat is indeed guaranteed to
lose points in the first play against a consistent
defector, but from round two onwards against such a
foe, it assumes the strategy itself, so the damage is
minimised.
In the initial competition, each program competed
against all others in a round-robin fashion; with the
twist that at some point it would play against a copy
of itself. With sufficiently long iterations, this allows
Tit-for-Tat to win even as the sole member of an
otherwise constantly defecting group: since when it
plays itself, it settles into a mutually beneficial
pattern of co-operation that brings more points per
round than mutual defection and hence counteracts
the few points lost in the opening play against

defectors. So whilst the defectors never give away
points, they miss opportunities to gain them,
opportunities which Tit-for-Tat seized to enable it to
climb the scoreboard. The crucial observation is that
an IPD tournament is a larger game still, in which
your objective is to get the highest possible total
score over a series of IPD games not to beat your
opponent in any given game.
To explain why Tit-for-Tat succeeds, Axelrod
advanced two key factors. The first he described as
niceness a nice program being one which would
never defect first. This has a massive advantage in
ensuring maximum points whenever two nice
programs encounter each other, as neither will move
away from mutual cooperation.
However, being too nice leads to exploitation,
either by actively malicious programs such as
always defect programs, or simply by random
behaviour. Being nice does not rule out the
possibility of defecting: it just requires a trigger
defection from the other player first. For Tit-For-Tat,
the trigger is a single defection: more complicated
triggers such as Tit-for-two-Tats are possible and in
fact this would have scored higher still against the
field of Axelrod's first tournament.
The second factor is forgiveness how long a
program punishes a defection for. Grim Trigger is
the ultimate grudge-bearing program: it initially
cooperates, but a single defection trigger will
convert it to always defect. Grim is technically a
nice program, and thus matches the performance of
Tit-for-Tat against other such strategies, whilst
typically scoring better against random behaviour.
However, against more complex systems that try to
gauge the friendliness of their opponent through
varying co-operation and defection, Grim scores
comparatively poorly.
Meta-gaming: beating Tit-for-Tat

through teamwork
Tit-for-Tat thus seemed to have an unassailable
combination of co-operating where possible, but
punishing where not. Yet in a twentieth-anniversary
competition run by computer scientist Graham
Kendall, with 223 entries, Tit-for-Tat couldn't claim
even one of the top three places. All of those
positions went instead to programs developed at
Southampton University, which were able to
recognise each other and then assumed master/slave
roles that maximised the payoff to the master

program at the expense of the slaves, which sunk
rapidly to the bottom of the performance table
through their heroic sacrifice.
Some see this as simply cheating, others as
ingenious meta-gaming beating the rules rather
than the opponents. Certainly it has limitations and
could fail spectacularly in rival configurations of the
Dilemma game; but with a bit of thought it becomes
apparent that Tit-for-Tat could also be accused of
gaming the system, and that the true point of running
Prisoner's Dilemma games is to find such insights.
The Southampton group was motivated by the
development of co-operative behaviour within multiagent systems, and tweaks to their approach should
shed light on the extent to which self-sacrificing
agents can help the system as a whole, or even just a
favoured group within it.
How then did the Southampton team out-play this
decades-old strategy? Their programs made use of
what makes the dilemma a dilemma in the first place
you can score even more than you would for mutual
co-operation, if only you could get the other player
to co-operate whilst you defect. Kendall's rules
allowed for more than one program to be submitted,
so you could submit just such a help, a program
following the suicidal always co-operate method,
except that all the other always defect programs
would get to feed upon it too. The solution was to
have the programs use pre-determined sequences of
co-operation and defection to act as a signature. On
encountering an appropriate sequence, the programs
would assume their roles, with the slaves switching
to constant co-operation to offer up bonus points to
the always-defecting masters. If, however, the
trigger sequence wasn't received from the opponent,
the always-defect approach was activated, to
minimise points available to the rival. In this way,
the masters would rack up more points than Tit-forTat, which would often be stuck with the paltry
rewards of mutual defection.
The most interesting aspects of this approach are
the issues of just what density of colluders is
required in the population it turned out that there

were probably too many in the 2004 event and
whether it would be possible for other programs to
spoof or manipulate the trigger sequences needed. In
evolutionary games where code is mixed or
programs are eliminated, it is possible for the slaves
to be removed from the game, stranding the masters
with the potential of being exploited during their
now-useless setup phase, and then performing no
better than normal always defect programs after
those steps. Nor can the fate of the slaves be entirely
ignored, as their sacrifice could reduce the average
performance per colluding program to below that of
tit-for-tat, depending on the ratios present in the
population.
The Prisoner's Dilemma seems destined to provoke
new ideas for years to come. Whilst it may be too
simple a model for real-world scenarios or human
psychology, it nonetheless has insights to offer as it
and its variations capture the essence of competition,
co-operation and individual versus group interest,
ideas which find wide application across the social
sciences.
About the Author
Graeme Taylor
completed his
MMath at the
University of Bath,
a four year
undergraduate
Masters offering a
wide range of
topics from Pure
and Applied
Mathematics and
related disciplines.
He is now a tutor
and postgraduate student at the University of
Edinburgh, where his research interest is in
computational aspects of Number Theory and
Algebraic Geometry.
FURTHER READING
Effective choice in the prisoners dilemma. R. Axelrod in Journal of Conflict Resolution Vol. 24 No. 1,
pages 3-25, 1980.
Does studying economics inhibit cooperation? R. Frank, T. Gilovich and D. Regan in Journal of Economic
Perspectives, Vol. 7, No. 2, pages 159-171, 1993.
Game Theory A Critical Introduction. S.P. Hargreaves-Heap and Y. Varoufakis, Routledge, 1995.
Theory of Games and Economic Behaviour. J. Von Neumann and O. Morgenstern, Princeton University
Press, 1944.
Autumn 2007
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mathematics beyond the

imaginary
iSquared
magazine 29
Population viability analysis: determining

whether a species will stick around
ildlife conservation is big news at the

minute, thanks to the medias
obsession with climate change. We are
being told how global warming is damaging the
environment and putting thousands of endangered
species at risk. Scientists are predicting that in
upcoming years many of these species will be lost

forever, resigned to the history books. But its not
only recently that scientists have been interested in
determining the fate of threatened species. In fact,
scientists have been making these types of
predictions for the past thirty years, mainly due to
Autumn 2007
Image courtesy of U.S. Fish and Wildlife Service
the development of Population Viability Analysis

models, or PVA models.
A Population Viability Analysis model is a
simulation model which is used to predict the
probability that a population of a specific species
will become extinct within a given number of years.
In other words, a PVA model provides a quantitative

estimate of extinction risk; therefore it is a way of
measuring how threatened a certain species is. It
calculates this measure of extinction using
demographical data (e.g. birth and death rates),
which is obtained from field work. It is a stochastic
model, which is to say it incorporates uncertainty.

This is what makes it a realistic model. If it didnt
incorporate uncertainty, all the model would tell you
is that if the mean population growth rate was
positive, the population would grow and if it was
negative, it would decline towards extinction. True
in the short term maybe, but not entirely useful when
it comes to making longer term predictions.
The first PVA model is credited to Mark Schaffer,
who in 1978 proposed a model to assist with the
management of the grizzly bears in Yellowstone
National Park. His model incorporated random
variability, and was used to calculate extinction
possibilities for the bear. Schaffer used his model to
calculate a MVP, or Minimum Viable Population,
for the grizzlies in Yellowstone. An MVP is
essentially the smallest number of individuals in a
population that enables the population to persist. On
initial simulations, Schaffer proposed that the MVP
for the grizzlies was 35, although in 1981 he
suggested that this should be increased to 50 90
bears due to uncertainty in the original estimate.
None the less, in coming up with a solution to the
problem of grizzly bear management in Yellowstone,
Schaffer had shown the world the part that

mathematics could play in wildlife conservation.
Extinction factors
In 1981, Schaffer went further with his PVA model

and started thinking about what the main causes of
extinction were. He discovered that there are four
main factors that contribute to a populations demise.
Demographic stochasticity, which is variation that
occurs in the demographics of the population, is one
of these factors. In large populations, these random
variations in birth and death rates tend to even
themselves out over time, but in smaller populations
such variations can have severe consequences.
However, a much more important factor is
environmental stochasticity. Variations in birth and
death rates due to random environmental changes,
such as disease or predation, have a much greater
impact on the dynamics of a population than
demographic stochasticity. Unlike demographic
stochasticity, environmental stochasticity can have
serious repercussions for even large populations.
Another factor that Schaffer looked at
was genetic stochasticity. Random changes
in the genetic makeup of a population can
affect the populations ability to survive
and reproduce, and therefore diminish a
Demographic stochasticity
populations numbers. This can make a bad
variations in birth/death rates
demographic situation in a small population
much worse. The last factor,
Environmental stochasticity much,
catastrophes, is quite different to the
random environmental changes
previous three. Whilst the previous factors
e.g. disease, predation
affect a population over time, a catastrophe
can have quite a sudden effect. Events like
Genetic stochasticity
large fires not only cause sudden loss of life,
changes in genetic makeup
but they can also destroy a populations
habitat, seriously damaging the
Catastrophes
populations dynamics. An unpredictable
e.g. large fires, volcanic eruption
event like this can destroy even the largest
of populations.
CAUSES OF EXTINCTION:
Interestingly, Schaffer found that rather

than a single one of these factors causing a
populations extinction, it is brought about
by a combination of factors. In 1986 two
scientists, M.E. Gilpin and M.E. Soule,
took this even further. They developed a
class of model through which extinction
could be categorised in context of the
events that lead up to the event. They called
Autumn 2007
Environmental
Genetic
R-Vortex
F-Vortex
Smaller population and increased

variability have destabilising effect
Decreased population size leads

to inbreeding depression
D-Vortex
A-Vortex
Increased spatial distribution,

population becomes fragmented
Outbreeding results in poorly

adapted offspring
Paths to extinction: Gilpin and Soules extinction vortices

these models Extinction Vortices. They proposed
that the population of a species spiraled towards
extinction in one of four ways, two of which are
brought about by environmental factors, and two
which are brought about by genetic factors.
The first Vortex they described was named the
R-Vortex. This method of extinction comes about
when there is a disturbance, such as a disease, which
from large numbers, such as populations where

larger numbers provide a better defence against
predators. Therefore, if the population becomes
fragmented, the local extinction rates for these
pockets increase. This in turn leads to the population
becoming more broken up, until in some cases the
population becomes so fragmented that it becomes
extinct.
Gilpin and Soule also considered genetic factors
that could lead to a populations extinction. The FSchaffer showed the world the Vortex
begins with a decrease in population size
part that mathematics could
which leads to an increase in inbreeding depression.
Inbreeding leads to a loss of variety in the
play in wildlife conservation
populations gene pool and this makes the
results in the population size shrinking, while at the
population prone to extinction. For example if all the
same time increasing the variability in the
members of the population are similar genetically,
population. An example of this would be an illness
then a disease could come along to which the
which killed significantly more females than males
majority of the population is vulnerable. This could
in the population, thus skewing the male to female
seriously reduce the size of the population and lead
ratio. Such an event could make the population more to extinction.
vulnerable to additional disturbances which would
The A-Vortex also deals with genetic factors, but
lower the population further, and further increase
instead of inbreeding leading to the populations
variability. So going back to the example, an
demise it is outbreeding that can cause extinction.
unbalanced sex ratio is prone to becoming more
This typically affects populations where the
unbalanced and in some cases this could lead to
members are specialised at living in their habitat.
extinction if the number of females became critical. When members of this population mate with
The other vortex that deals with environmental
members of a different population, which has
factors was named the D-Vortex. This vortex
adapted to living in a different habitat, the offspring
begins with a decrease in population size, which
can be suited to neither habitat, and so will struggle
increases the variability and the spatial distribution
to survive. Therefore, this cross-breeding can lead to
of the population. This results in the population
extinction for the population.
becoming broken up and fragmented. This type of
The use of genetics in PVA has been questioned
vortex would affect a population which benefits
by some. In 1988, Russell Lande suggested that
Image courtesy of U.S. Fish and Wildlife Service
demographic factors, such as those which influence

population growth, were much more important than
genetic factors in determining the chance that a
species would become extinct. He put forward that
by the time a population is facing risk of extinction
from genetic factors, the population is already
demographically close to its demise. Therefore, he
suggested that it was unnecessary to include genetic
factors in a PVA model.
Sensitivity analysis of PVA models helped

conservationists who were trying to protect
the loggerhead sea turtle
A tool for turtles

It was in the United States where PVA began to gain
popularity. Ecologists and federal agencies, acting in
accordance with the Endangered Species Act of
1966 and the National Forest Management Act of
1976, needed methods to evaluate extinction risk,
and they found that PVA met their needs. Of course,
as with all mathematical models of the time, it was
in the late 80s and early 90s when the use of PVA
really exploded, with the aid of more advanced
computers.
The purpose of early PVA models, such as the one
created by Schaffer, was to determine the MVP of
the population, but as it became understood that
populations become increasingly vulnerable to
extinction as they become smaller, emphasis soon
shifted to other uses. PVA started to be more
commonly used in aiding conservation management
decisions, as the major advances in computing
software at the time made it a lot easier to predict the
results of proposed methods. Scientists were also
beginning to use PVA to perform a sensitivity
analysis on a population, which enabled them to see
which area of conservation it would be most
beneficial to focus on.
A sensitivity analysis is where a variable in the
PVA model is taken and studied to see how much it
affects the population of the species that is being
looked at. This is done by changing the value of the
variable, whilst keeping other variables the same,
and running a simulation of the model. The value of
some measure of extinction risk is then recorded.
Many simulations are run, with each taking a
different value for the variable that is being looked at,
and the measures of extinction risk are compared. If
the measures vary greatly, then the variable in
question is found to have a large effect on the
population, and conservation efforts should be
focused on trying to keep that variable at an
optimum value.
The example of the loggerhead sea turtle shows
how useful this process can be. For years,
conservation efforts for the turtle had focused on
ensuring that eggs on nesting beaches were
undisturbed, so that they were able to hatch, and the
turtles could safely enter the sea. A sensitivity
analysis was carried out and found that survivorship
through the egg stage actually had little effect on
population outcomes. It was in fact, survivorship
through the oceanic juvenile stage which was critical
to the health of the population. Therefore, in light of
this startling discovery, conservation efforts were
shifted away from egg survival and towards juvenile
survivorship. This was achieved by focusing on
incorporating turtle-excluder devices into fishing
nets.
Despite these benefits of using PVA, many people
question whether it is much use in conservation
biology at all. The ability of PVA to accurately
Autumn 2007
assess extinction risks is forever being called into

question. It obviously only offers predictions, but
just how good are these predictions?
The main problem with PVA is that it requires a
large quantity of field data. This is often difficult or
impossible to acquire, especially as PVA mostly
deals with endangered species where data is
extremely limited. So, as the predictive power of
PVA diminishes with smaller data sets, many have
said that it has little value in predicting extinction
risks for endangered species. Even when an adequate
data set is available, it is possible that a PVA can
still produce large errors when making extinction
rate predictions. But while some are calling for PVA
to be replaced by other methods, many argue that it
remains the best tool available for estimates of
extinction risk. They especially point out how useful
sensitivity analysis can be in deducing the most effective way of managing conservation, as in the example of the loggerhead sea turtle. Many scientists wish
for a fixed definition of PVA to be accepted by the
scientific community. They also want to introduce
scientific standards of quality by which all PVA are
judged. However, this could be easier said than done.
Either way, in the future many improvements will
be made to PVA. As with any model, refinements
are constantly being made in order to improve its
reliability, but scientists are also aiming to broaden it.

They are working on developing a reliable PVA for
plants, and are also working on incorporating recent
genetic advances into the model. (Lande has now
acknowledged the role that genetics play in extinction risk).
Exactly how accurate a predicting tool PVA
can become is hard to predict. At this
moment in time though, PVA is a very useful tool
in wildlife conservation, whether it is in identifying
populations at risk, or helping to forecast the
effectiveness of management plans. Just ask the
loggerhead sea turtle.
About the Author
Daniel Rowe was
born in Somerset,
where he lived until
he went to the
University of Bath to
study mathematics.
While at Bath he
developed a
particular interest in
mathematical biology.
Daniel recently graduated and is currently
actively seeking employment.
FURTHER READING
Minimum population sizes for species conservation. Mark L. Shaffer in BioScience, Vol. 31, No. 2,
pages 131-134, 1981.
Minimum viable populations: Processes of species extinction. M. E. Gilpin and M. E. Soule, pages 19-34 in
Conservation biology: the science of scarcity and diversity, edited by M.E. Soule, Sinauer Associates, 1986.
A stage-based population model for loggerhead sea turtles and implications for conservation.
Deborah T. Crouse, Larry B. Crowder and Hal Caswell in Ecology, Vol. 68, No. 5, pages 1412-1423, 1987.
Precision of Population Viability Analysis. Stephen P Ellner, John Fieberg, Donald Ludwig and Chris Wilcox in
Conservation Biology, Vol. 16, No. 1, pages 258261, 2002.
Sikaku
Killer Sudoku
Bridges
Solutions to puzzles on pages 37-38:

puzzles
Prize crossword
1
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
Across
1 Father of set theory not arc (anag) (6)
7 Except on a null set - describes region where
some property holds true (abbr) (2)
9 Specific mode of vibration march ion (anag) (8)
10 Root of a polynomial (4)
11 Mathematical function related to exponential (2)
12 Nineteenth century French mathematician (6)
14 In trigonometry, inverse of tangent (abbr) (3)
16 Type of coordinate system (5)
17 Unit of power (4)
20 Discrete quantity physical theory involving
wave-particle duality (7)
21 Obvious (7)
24 Quantity (5)
25 Mathematical approximation of reality (5)
26 Type of equation involving derivatives with
respect to several variables (abbr) (3)
29 Irish mathematician/physicist theorem in vector
analysis relating surface and line integrals (6)
31 Common abbreviation used in mathematical
reasoning (2)
32 Fraction between zero and one (4)
33 Branch of mathematics concerned with
space extension of geometry (8)
34 14th letter in the Greek alphabet (2)
Win a copy of the Concise

Oxford Dictionary of
Mathematics.
Send your completed entry by
31st October to: Prize Crossword,
iSquared magazine, 1 Pound
Cottages, Shillinglee, Godalming,
Surrey GU8 4SZ. The first correct
entry drawn
after the
closing date
will win a
dictionary.
Only one
entry per
person is
permitted.
The solution
and winner
will be printed in the
next issue.
35 Property of two integers depending on the sign

of both numbers (6)
Down
1 Recurring at regular intervals type of group (6)
2 Types of graph in discrete mathematics
branched objects (5)
4 Object with a non-integer dimension (6)
5 Single-holed surface of revolution (5)
6 Electromagnetic radiation (5,4)
8 Force applied against inertia (6)
13 Fixed point of a conic section (5)
18 Method of mathematical proof (9)
19 Degree size, often in group theory (5)
22 Logic operation (3)
23 Force in roots (anag) (7)
24 Point of intersection of two sides of a polygon (6)
27 Set containing subsets discrete unit
tin tye (anag) (6)
28 Prolific Swiss mathematician who gave his
name to a numerical method for solving
differential equations (5)
30 Entwined circle topological theory (4)
32 Side of triangle in trigonometry (abbr) (3)
Autumn 2007
Sikaku
How to Play
Divide the grid into rectangles, so
that each rectangle contains one
digit and exactly the number of
squares indicated by that digit. For
example, a rectangle containing
the number 2 must contain two
squares of the grid.
Solution on page 35.
Puzzle supplied by Clarity Media.
Killer Sudoku
How to Play
As for regular sudoku, fill in
the grid so that every row,
column and 3x3 group of
cells contain the numbers 1
to 9. However, the numbers
in the cells enclosed within
each cage (dotted lines
denote cages) must add up
to the total given for that
cage. No cage can contain
the same number twice.
Puzzle supplied by
Clarity Media.
puzzles
Bridges
How to Play
By joining the islands (encircled numbers) with straight lines, create a continuous path
between all the islands. The circled numbers indicate how many bridges must join that island.
Up to two bridges can be created between any two islands. Bridges can only exist between
islands that are directly adjacent in either the north, south, east or west directions.
Puzzle supplied by Clarity Media.
Autumn 2007
There is no branch of mathematics, however abstract, which

may not someday be applied to the phenomena of the real world.
Nicolai Lobachevsky
Nature's great book is written

in mathematical symbols
Galileo
xkcd
By
Randall
Munroe
Visit our website
isquaredmagazine.co.uk
www.
Autumn 2007

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isquared

mathematics beyond the imaginary

By Catherine Buchanan and David Stern

Cover Story: Can mathematics solve

The mathematics of being nice

Any Rubiks cube can be solved in a maximum of 26 moves

simplified the model and

Sand wave dynamics are

mathematical model for sand

and the design of offshore

in defining the shape

biological evidence of tumour

Instability can have

for the planned high-voltage cable

Optimal control theory is a

Image courtesy of Dr Timothy Triche/National Cancer Institute

resources is a prominent problem

Complicite, explores the

By Catherine Buchanan and David Stern

he omnipresence of mathematics in our

previous decisions. This is best illustrated by the

Here, the distances between intersections are marked

intersection H. First, f(X) is defined to be the

This method of finding the best path in a step by step

Building the sailing model

f(C) = f(B) + 9 = 5 + 9 = 14.

To formulate a real-life problem mathematically a

f(E) = f(D) + 3 = 8 + 3 = 11,

right was found by practical experimentation. Mike

Dynamic programming for the

take a limited number of specified values between

The tacking penalty chosen is to halve the boat's

them. Part of the challenge of creating a dynamic

Sailing models using a computer

Even after eliminating some positions the problem

The eastern bank of the lake is reached at this stage

Sailing in a sinusoidal wind:

The wind has a constant strength of 35. An optimal

The boats would take similar routes, travelling in a

Born in Budapest in 1903, John

Image courtesy of Los Alamos National Laboratory

If people do not believe that mathematics is simple, it is only because

of the newly formed department

Neumann architecture and is used

long with HIV/AIDS and tuberculosis,

Images courtesy of CDC (Centers for Disease Control and Prevention)

How mathematical models

Top: a female Anopheles mosquito feeds on a human host

grow inside red blood cells, inevitably leading to the

Blood cell models

Newly created cells

that having a stronger parasite has on the outcome of

Life cycle of red blood cells in a malaria patient

Unparasitised cells become

Free parasites die within 20

this number is between 8 and 32). The free parasites

The majority of deaths from malaria occur

Map image courtesy of CIA Factbook

Whilst this is an interesting model which can shed

enough to generate rapid parasite

In general there is a balance to be achieved

Mathematical modelling provides

fitted a slightly different model of a malaria infection