Applications of computer modelling to behavioural coordination
Ph.D. Thesis

©A R Ludlow B.Sc., Ph.D.

September 5, 2004
Home page:
Home page:

Instructions for downloading

To my surprise, I discovered that my paper The behaviour of a model animal (Ludlow, 1976) is still being cited quite regularly. There are other papers and a Ph.D. thesis on the ‘Model Animal’ [1, 2, 3, 4, 5, 6] The fullest and most recent account is the thesis (1983), so I have scanned it in and made it available as a .pdf file.

Chapter 5 is now included (5th Sep 2004) Chapters 1-4 cover all the work on the ‘Model Animal’ [2]. Chapter 5 covers modelling moth orientation.

The thesis [6] has been scanned into the computer using optical character recognition, followed by extensive proof reading. There are bound to be some discrepancies that have been overlooked but I have been careful to make no deliberate changes to the text which was completed in November 1983.

The following bullet points allow modern comments on the original text:

Chapter 1
is a philosophical discussion of the purpose of modelling. It seems dated now but was necessary in 1983 because theoretical studies in behaviour were not widely appreciated in those days and a theoretical Ph.D. had to be justified.
Chapter 2
describes the ‘Model Animal’ as it appeared in [2]. The text is a review (in 1983) of what I wished I had said in 1976, together with a few changes in the program.
Chapter 3
is ‘The consequences of inhibition’. It covers the original ‘winner takes all’ model and shows how this can be generalised to suggest mechanisms of pattern recognition and even learning. Much of this material was not published because of a career change when I became head of biological modelling at the Forestry Commission.
Chapter 4
is ‘The consequences of fatigue’ and discusses processes which may lead to an alternation of activities. Great stress is laid on how these processes can be identified from analying changes in bout length.
Chapter 5
is ‘Mechanisms of Moth Orientation’. It describes our work on the mechanisms by which male moths find a female emitting sex-pheromone. The emphasis is on modelling the visual inputs and asks whether moths had beaten Pythagoras to his theorem. Essentially the same ground is summariseded in one of my lecture notes:

The thesis may be downloaded by clicking on: The file is 1 MB and the latest version involved some corrections of the optical character recognition were made on 5th September 2004.

In addition, the reader may wish to consult three other papers bound into the back of the thesis. [2, 4, 5]


[1]     A.R. Ludlow. Reciprocal inhibition and a naive model of behaviour. Postgraduate conference on animal behaviour, Cambridge, 1972, 1972.

[2]     A.R. Ludlow. The behaviour of a model animal. Behaviour, 58:131-172, 1976.

[3]     A.R. Ludlow. The importance of temporal coupling between feeding and drinking -- simulations prompted by Toates’paper. Behavioural and Brain Sciences, 2:110-111, 1979.

[4]     A.R. Ludlow. The evolution and simulation of a decision maker. In F.M. Toates and T.R. Halliday, editors, Analysis of motivational processes, pages 273-296. Academic Press, 1980.

[5]     A.R. Ludlow. Towards a theory of thresholds. Animal Behaviour, 30:253-267, 1982.

[6]     A.R. Ludlow. Applications of computer modelling to behavioural coordination. PhD thesis, University of London, 1983.