mathematics beyond the imaginary
Autumn 2007

Extinction modelling Prisoner’s Dilemma John von Neumann





Volume 1 Number 1

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.


Sarah Shepherd
Cover image by Eric Isselée iSquared magazine, published quarterly. Issue 1 Number 1 (ISSN 1755-7275). Postal address: iSquared magazine, 1 Pound Cottages, Shillinglee, Godalming, Surrey, GU8 4SZ. Email: Website: 2 iSquared magazine
Printed in the UK by THE MAGAZINE PRINTING COMPANY Autumn 2007

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.

autumn 2007


Cover Story: Can mathematics solve the problem of malaria?
By Deborah Cromer How mathematical models can shed light on one of the world’s deadliest diseases.


The mathematics of being nice
By Graeme Taylor Mutual co-operation gives the best outcome, but what is the winning strategy in game theory’s Prisoner’s Dilemma?


Survival or extinction?
By Daniel Rowe Population viability analysis: determining whether a species will stick around.


News Mathematical greats
The life and work of the Hungarian-born US mathematician, John von Neumann.

4 14 22 29 36 39
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Book review
Symmetry and the Monster, the story of one of the greatest quests of mathematics.

Subscription form Puzzles Endnotes

Rubik’s number
The Rubik’s 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 Rubik’s cube is known as “God’s 4 iSquared magazine

Any Rubik’s 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 God’s number is in the “low 20’s”. 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

for the planned high-voltage cable between England and the Netherlands.

Möbius puzzle
A 75-year old problem involving the Möbius strip has finally been solved, by two researchers at University College, London. The Möbius 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 Möbius 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.

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.

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

The Möbius strip has been described mathematically using differential equations

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David Benbennick

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

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

A Disappearing Number

Joris Jan-Bos

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 Ramanujan’s footsteps, and her lover, who follows after her death. The importance of Ramanujan’s 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

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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

By Catherine Buchanan and David Stern

Image by Manfred E. Fritsche (licensed by Creative Commons Attribution ShareAlike 2.0 Germany License)

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.




3 2 3









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

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.

f(A) = 0, f(B) = 5,

Building the sailing model

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}

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

= 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:

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.

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 (θ).













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
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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 θ 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.

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.

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 (θ) is restricted, so that θ is allowed to

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.

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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. 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 Vanderbei’s 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.

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.

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 12 iSquared magazine

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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 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.

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.

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,, 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.

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mathematical greats
The life and work of the Hungarian-born US mathematician, John von Neumann If people do not believe that mathematics is simple, it is only because they do not realise how complicated life is. 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 1930’s, after Hitler’s 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 14 iSquared magazine

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

Image courtesy of Los Alamos National Laboratory

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 Kubrick’s 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 iSquared magazine 15

Can mathematics solve the problem of

How mathematical models can shed light on one of the world’s deadliest diseases
long with HIV/AIDS and tuberculosis, malaria is one of the World Heath Organisation’s “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 world’s population is exposed to malaria and between 300–660 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 world’s 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 16 iSquared magazine

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 patient’s 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)

Top: a female Anopheles mosquito feeds on a human host Bottom: scanning electron micrograph (SEM) of an Anopheles mosquito

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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

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 β. This means that parasitised cells are created at a rate of βu(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

Newly created cells replace aging cells

Life cycle of red blood cells in a malaria patient

Unparasitised cells u(t)

Unparasitised cells become parasitised by combining with a free parasite

Parasitised cells p(t)

Unparasitised cells die after 120 days due to ageing

Rupture of parasitised cells creates 8–32 new free parasites

Free parasites m(t)

Free parasites die within 20 minutes, if they do not invade a red blood cell

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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.

Beta paradox
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

The majority of deaths from malaria occur in Africa children below the age of five

The worldwide distribution of malaria: dark colouring indicates high risk regions

iSquared magazine 19

Map image courtesy of CIA Factbook


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 patient’s 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. 20 iSquared magazine
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

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 WHO’s ‘Big 3’ will be reduced to the ‘Big 2’.●

“Mathematical modelling provides a safe and inexpensive way to explore how the infection behaves”

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.

iSquared magazine 21

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 Évariste 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 Galois’s 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 22 iSquared magazine 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, Lie’s 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

Émile 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 “Janko’s 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 store…it 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 1970’s 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 Lie’s 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 iSquared magazine 23

building trust in the Iterated Prisoner’s Dilemma

Mutual co-operation gives the best outcome, but what is the winning strategy in game theory’s Prisoner’s 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 Prisoner’s 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.

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The possible outcomes of the dilemma are given in the following table: 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: Prisoner A defects

Prisoner B defects
Each serves 4 years

Prisoner B co-operates
Prisoner A serves 1 year, Prisoner B serves 5 years Each serves 2 years

Prisoner A serves 5 years, Prisoner B serves 1 year


(1,1) (4,0) (0,4) (3,3)


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 iSquared magazine 25

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 other’s 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 26 iSquared magazine
Autumn 2007

Tit-for-Tat strategy is: · In round 1, co-operate. · In round n for n ≥ 2, play your opponent's strategy from round n–1. 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 iSquared magazine 27

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 FURTHER READING

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.

Effective choice in the prisoner’s 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.

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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 media’s 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 30 iSquared magazine


upcoming years many of these species will be lost forever, resigned to the history books. But it’s 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 iSquared magazine 31

model, which is to say it incorporates uncertainty. This is what makes it a realistic model. If it didn’t 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 population’s 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 population’s ability to survive and reproduce, and therefore diminish a Demographic stochasticity population’s 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 population’s habitat, seriously damaging the Catastrophes population’s dynamics. An unpredictable e.g. large fires, volcanic eruption event like this can destroy even the largest of populations. Interestingly, Schaffer found that rather than a single one of these factors causing a population’s 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

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Smaller population and increased variability have destabilising effect

Decreased population size leads to inbreeding depression

Increased spatial distribution, population becomes fragmented

Outbreeding results in poorly adapted offspring

Paths to extinction: Gilpin and Soule’s 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 population’s extinction. The ‘F“Schaffer 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” population’s 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 population’s 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 iSquared magazine 33

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 80’s and early 90’s 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

Image courtesy of U.S. Fish and Wildlife Service

34 iSquared magazine

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 FURTHER READING

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.

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 258–261, 2002.

Solutions to puzzles on pages 37-38:

iSquared magazine 35


Killer Sudoku


Prize crossword
1 2 3 9 10 11 13 14 15 18 20 23 24 25 28 29 30 32 33 34 35 31 26 27 21 16 19 22 17 12 4 5 6 7 8

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.

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)

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) 3 17th letter in the Greek alphabet (3) 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) 15 19th letter in the Greek alphabet (3) 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

36 iSquared magazine

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. Solution on page 35.

Puzzle supplied by Clarity Media.

iSquared magazine 37


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. Solution on page 35. Puzzle supplied by Clarity Media.

38 iSquared magazine

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


By Randall Munroe

iSquared magazine 39

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