The Geometry of Stimulus Control

Ghirlanda, Stefano and Enquist, Magnus (1999) The Geometry of Stimulus Control. [Journal (Paginated)]

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Many studies, both in ethology and comparative psychology, have shown that animals react to modifications of familiar stimuli. This phenomenon is often referred to as generalisation. Most modifications lead to a decrease in responding, but to certain new stimuli an increase in responding is observed. This holds for both innate and learned behaviour. Here we propose a heuristic approach to stimulus control, or stimulus selection, with the aim of explaining these phenomena. The model has two key elements. First, we choose the receptor level as the fundamental stimulus space. Each stimulus is represented as the pattern of activation it induces in sense organs. Second, in this space we introduce a simple measure of `similarity' between stimuli by calculating how activation patterns overlap. The main advantage we recognise in this approach is that the generalisation of acquired responses emerges from a few simple principles which are grounded in the recognition of how animals actually perceive stimuli. Many traditional problems that face theories of stimulus control (e.g. the Spence-Hull theory of gradient interaction or ethological theories of stimulus summation) do not arise in the present framework. These problems include the amount of generalisation along different dimensions, peak-shift phenomena (with respect to both positive and negative shifts), intensity generalisation, and generalisation after conditioning on two positive stimuli.

Item Type:Journal (Paginated)
Keywords:generalization, stimulus control, evolution, neural networks
Subjects:Biology > Animal Behavior
Biology > Theoretical Biology
Neuroscience > Computational Neuroscience
Computer Science > Neural Nets
ID Code:877
Deposited By:Ghirlanda, Dr. Stefano
Deposited On:18 Jul 2000
Last Modified:11 Mar 2011 08:54

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References in Article

Select the SEEK icon to attempt to find the referenced article. If it does not appear to be in cogprints you will be forwarded to the paracite service. Poorly formated references will probably not work.

[Atkinson & Estes 1963]

Atkinson, R. R. & Estes, W. K. 1963. Stimulus sampling theory. In Handbook of Mathematical Psychology (Ed. by R. D.

Luce & R. B. Bush), pp. 121-268. New York: Wiley.

[Baerends 1975]

Baerends, G. 1975. An evaluation of the conflict hypothesis as an explanatory principle for the evolution of displays. In

Function and evolution of behaviour (Ed. by G. Baerends, C. Beer & A. Manning), pp. 187-227. Oxford: Clarendon Press.

[Baerends & Drent 1982]

Baerends, G. P. & Drent, R. H. 1982. The herring gull and its egg. Behaviour, 82, 1-416.

[Baerends & Krujit 1973]

Baerends, G. P. & Krujit, J. P. 1973. Stimulus selection. In Constraints on Learning (Ed. by R. A. Hinde &

J. Stevenson-Hinde), pp. 23-50. New York: Academic Press.

[Baron 1973]

Baron, A. 1973. Postdiscrimination gradients of human subjects on a tone continuum. Journal of Experimental Psychology,

101, 337-342.

[Blough 1975]

Blough, D. S. 1975. Steady state data and a quantitative model of operant generalization and discrimination. Journal of

Experimental Psychology: Animal Behavior Processes, 104, 3-21.

[Brennan & Riccio 1973]

Brennan, J. F. & Riccio, D. C. 1973. Stimulus control of avoidance behavior in rats following differential or nondifferential

pavlovian training along dimensions of the conditioned stimulus. Journal of Comparative and Physiological Psychology, 85,


[Coren & Ward 1989]

Coren, S. & Ward, L. M. 1989. Sensation and perception. Fort Worth, TX: Harcourt Brace Jovanovitch, Publishers.

[Galizio 1980]

Galizio, M. 1980. Conditional gradient displacements: the effects of conditional discrimination training on human auditory

frequency generalization. Journal of Experimental Psychology: Human Learning and Memory, 6, 766-777.

[Galizio & Baron 1976]

Galizio, M. & Baron, A. 1976. Label training and auditory generalization. Learning and Motivation, 7, 591-602.

[Ghirlanda & Enquist 1999]

Ghirlanda, S. & Enquist, M. 1999. Artificial neural networks as models of stimulus control. Animal Behaviour, 56,


[Guttman 1965]

Guttman, N. 1965. Effects of discrimination formation on generalization from the positive-rate baseline. In Stimulus

Generalization (Ed. by D. I. Mostofsky), pp. 211-217. Stanford, CA: Stanford University Press.

[Guttman & Kalish 1956]

Guttman, N. & Kalish, H. I. 1956. Discriminability and stimulus generalization. Journal of Experimental Psychology, 51,


[Hanson 1959]

Hanson, H. M. 1959. Effects of discrimination training on stimulus generalization. Journal of Experimental Psychology, 58,


[Hearst 1968]

Hearst, E. 1968. Discrimination training as the summation of excitation and inhibition. Science, 162, 1303-1306.

[Hinde 1970]

Hinde, R. A. 1970. Animal Behaviour. Tokyo: McGraw-Hill Kogakusha, Ltd., 2nd edition.

[Huff et al.Huff, Sherman & Cohn 1975]

Huff, R. C., Sherman, J. E. & Cohn, M. 1975. Some effects of response-independent reinforcement in auditory

generalization gradients. Journal of the Experimental Analysis of Behavior, 23, 81-86.

[Hull 1943]

Hull, C. L. 1943. Principles of Behaviour. New York: Appleton-Century-Crofts.

[Jenkins & Harrison 1960]

Jenkins, H. M. & Harrison, R. H. 1960. Effect of discrimination training on auditory generalization. Journal of Experimental

Psychology, 59, 246-253.

[Jenkins & Harrison 1962]

Jenkins, H. M. & Harrison, R. H. 1962. Generalization gradients of inhibition following auditory discrimination training.

Journal of the Experimental Analysis of Behavior, 5, 435-441.

[Kalish & Guttman 1957]

Kalish, H. I. & Guttman, N. 1957. Stimulus generalization after equal training on two stimuli. Journal of Experimental

Psychology, 53, 139-144.

[Kalish & Guttman 1959]

Kalish, H. I. & Guttman, N. 1959. Stimulus generalization after training on three stimuli: a test of the summation

hypothesis. Journal of Experimental Psychology, 57, 268-272.

[Koehler & Zagarus 1937]

Koehler, O. & Zagarus, A. 1937. Beiträge zum Brutverhalten des Halsbandregepfeifers (Charadrius h. hiaticula L.).

Beiträge zur Fortpflanzungsbiologie der Vögel, 13, 1-9.

[Mackintosh 1974]

Mackintosh, N. J. 1974. The psychology of animal learning. London: Academic Press.

[Marsh 1972]

Marsh, G. 1972. Prediction of the peak shift in pigeons from gradients of excitation and inhibition. Journal of Comparative

and Physiological Psychology, 81, 262-266.

[Nosofsky 1987]

Nosofsky, R. M. 1987. Attention and learning processes in the identification and categorization of integral stimuli. Journal

of Experimental Psychology, 13, 87-108.

[Ohinata 1978]

Ohinata, S. 1978. Postdiscrimination shift of the goldfish (Carassius Auratus) on a visual wavelength continuum. The

Annual of Animal Psychology, 28, 113-122.

[Pearce 1994]

Pearce, J. M. 1994. Similarity and discrimination: a selective review and a connectionist model. Psychological Review, 101,


[Purtle 1973]

Purtle, R. B. 1973. Peak shift: a review. Psychological Bullettin, 80, 408-421.

[Razran 1949]

Razran, G. 1949. Stimulus generalization of conditioned responses. Psychological Bullettin, 46, 337-365.

[Rescorla 1976]

Rescorla, R. A. 1976. Stimulus generalization: some predictions from a model of pavlovian conditioning. Journal of

Experimental Psychology, 2, 88-96.

[Rilling 1977]

Rilling, M. 1977. Stimulus control and inhibitory processes. In Handbook of operant behaviour, pp. 432-480.

[Shanks 1995]

Shanks, D. S. 1995. The psychology of associative learning. Cambridge: Cambridge University Press.

[Shepard 1987]

Shepard, R. N. 1987. Toward a universal law of generalization for psychological science. Science, 237, 1317-1323.

[Spence 1937]

Spence, K. W. 1937. The differential response in animals to stimuli varying in a single dimension. Psychological Review, 44,


[Stevenson 1966]

Stevenson, J. G. 1966. Stimulus generalization: the ordering and spacing of test stimuli. Journal of the Experimental Analysis

of Behavior, 9, 457-468.

[Thomas & Setzer 1972]

Thomas, D. R. & Setzer, J. 1972. Stimulus generalization gradients for auditory intensity in rats and guinea pigs.

Psychonomic Science, 28, 22-24.

[Tinbergen 1942]

Tinbergen, N. 1942. An objective study of the innate behaviour of animals. Bibliotheca biotheorica, 1, 39-98.

[van Kampen & de Vos 1995]

van Kampen, H. S. & de Vos, G. J. 1995. A study of blocking and overshadowing in filial imprinting. Quarterly Journal of

Experimental Psychology, 49B, 346-356.

[Vos 1995]

Vos, D. R. 1995. The development of sex recognition in the zebra finch. PhD thesis, Rijksuniversiteit Groningen, The


[Zielinski & Jakubowska 1977]

Zielinski, K. & Jakubowska, E. 1977. Auditory intensity generalization after CER differentiation training. Acta

Neurobiologiae Experimentalis, 37, 191-205.


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