Abstract
One of the main challenges to the adaptionist program in general and the use of optimization models in behavioral and evolutionary ecology, in particular, is that organisms are so constrained' by ontogeny and phylogeny that they may not be able to attain optimal solutions, however those are defined. This paper responds to the challenge through the comparison of optimality and neural network models for the behavior of an individual polychaete worm. The evolutionary optimization model is used to compute behaviors (movement in and out of a tube) that maximize a measure of Darwinian fitness based on individual survival and reproduction. The neural network involves motor, sensory, energetic reserve and clock neuronal groups. Ontogeny of the neural network is the change of connections of a single individual in response to its experiences in the environment. Evolution of the neural network is the natural selection of initial values of connections between groups and learning rules for changing connections. Taken together, these can be viewed as “design parameters”. The best neural networks have fitnesses between 85% and 99% of the fitness of the evolutionary optimization model. More complicated models for polychaete worms are discussed. Formulation of a neural network model for host acceptance decisions by tephritid fruit flies leads to predictions about the neurobiology of the flies. The general conclusion is that neural networks appear to be sufficiently rich and plastic that even weak evolution of design parameters may be sufficient for organisms to achieve behaviors that give fitnesses close to the evolutionary optimal fitness, particularly if the behaviors are relatively simple.
This is a preview of subscription content, log in via an institution to check access.
Similar content being viewed by others
References
Aarts, E., Korst, J.: Simulated annealing and Boltzmann machines. New York: Wiley 1989
Aoki, C., Siekevitz, P.: Plasticity in brain development. Sci. Am. 259, 56–67 (1988)
Averill, A. L., Prokopy, R. J.: Residual activity of oviposition deterring pheromone in Rhagoletis pomonella (Diptera: Tephritidae) and female response to infested fruit. J. Chem. Ecol. 13, 167–177 (1987)
Bounds, D. G.: New optimization methods from physics and biology. Nature 329, 215–219 (1987)
Brady, R. M.: Optimization strategies gleaned from biological evolution. Nature 317, 804–806 (1985)
Charnov, E. L.: Phenotypic evolution under Fisher's fundamental theorem of natural selection. Heredity, 62, 113–116 (1989)
Cowan, J. D., Sharp, D. H.: Neural networks. Quart. Rev. Biophys. 21, 365–427 (1988)
Crnjar, R. M., Prokopy, R. J.: Morphological and electrophysiological mapping of tarsal chemoreceptors of oviposition-deterring pheromone in Rhagoletis pomonella flies. J. Insect Physiol. 28, 393–400 (1982)
Dehaene, S., Changeux, J.-P., Nadal, J.-P.: Neural networks that learn temporal sequences by selection. Proceedings of the National Academy of Sciences 84, 2727–2731 (1987)
Edelman, G.: Neural Darwinism. New York: Basic Books 1987
Edwards, D. H., Mulloney, B.: Synaptic integration in excitatory and inhibitory crayfish motoneurons. J. Neurophysiol. 57, 1425–1445 (1987)
Evans, S. M.: Non-associative behavioural modifications in nereid polychaetes. Nature 211, 945–948 (1986)
Evans, S. M. Behavior in polychaetes. Quart. Rev. Biol. 46, 379–405 (1971)
Evans, S. M.: Effects of changes in sensory input on patterns of tubicolous activity in the polychaete Platynereis dumerilii. Marine Behav. Physiol. 7, 307–319 (1981)
Evans, S. M., Downie, P. J.: Decision-making processes in the polychaete Platynereis dumerilii. Anim. Behav. 34, 472–479 (1986)
Finkel, L. H., Edelman, G. M.: Interactions of synaptic modification rules within populations of neurons. Proc. Nat. Acad. Sci. USA 82, 1291–1295 (1985)
Friesen, W. O., Wyman, R. J.: Analysis of Drosophila motor neuron activity patterns with neural analogs. Biol. Cybernetics 38, 41–50 (1980)
Getting, P. A.: Mechanisms of pattern generation underlying swimming in Tritonia. II. Network reconstruction. J. Physiol. 49, 1017–1035 (1983a)
Getting, P. A.: Mechanisms of pattern generation underlying swimming in Tritonia. III. Intrinsic and synaptic mechanisms for delayed excitation. J. Physiol. 49, 1036–1050 (1983b)
Getting, P. A.: Comparative analysis of invertebrate central pattern generators. In: Cohen, A. (ed.) Neural control of rhythmic movements in vertebrates, chap. 4, pp. 101–127. New York: Wiley 1988
Getz, W. M.: A neural network for processing olfactory-like stimuli. Biol. Cybernetworkics, in press
Gould, S. J., Lewontin, R. C.: The spandrels of San Marco and the Panglossian paradigm: a critique of the adpationist program. Proc. Rl. Soc Lond. B 205, 581–598 (1979)
Grafen, A.: On the uses of data on lifetime reproductive success. In: Clutton-Brock, T. H. (ed.) Reproductive success, pp. 454–471. Chicago: University of Chicago Press 1988
Hinton, G. E., Nowlan, S. J.: How learning can guide evolution. Complex Systems 1, 495–502 (1987)
Hoffmann, G. W., Benson, M. W., Bree, G. M., Kinahan, P. E.: A teachable neural network based on an unorthodox neuron. Physica 22D, 233–246 (1986)
Kauffman, S., Levin, S.: Towards a general theory of adaptive walks on rugged landscapes. J. Theor. Biol. 128, 11–45 (1987)
McClelland, J. L., Rumelhart, D. E., PDP Research Group: Parallel distributed processing. Volume 2: Psychological and biological models. Cambridge, MA: MIT Press 1986
Mangel, M.: Oviposition site selection and clutch size in insects. J. Math. Biol. 25, 1–22 (1987)
Mangel, M.: The evolution of optimal behavior by natural selection. Preprint, Department of Zoology, University of California, Davis
Mangel, M., Clark, C. W.: Dynamic modeling in behavioral ecology. Princeton, NJ: Princeton University Press 1988
Maynard Smith, J.: When learning guides evolution. Nature 329, 761–762 (1987)
Mulloney, B., Perkel, D. H.: The roles of synthetic models in the study of central pattern generators. In: Cohen, A. (ed.) Neural control of rhythmic movements in vertebrates, Chap. 11, pp. 415–453. New York: Wiley 1988
Miller, J. R., Strickler, K. L.: Finding and accepting host plants. In: Bell, W. J., Carde, R. T. (eds.) Chemical ecology of insects. London: Chapman and Hall 1984
Papaj, D. R., Prokopy, R. J.: Phytochemical basis of learning in Rhagoletis pomonella and other herbivorous insects. J. Chem. Ecol. 12, 1125–1143 (1986)
Prokopy, R. J., Averilli, A. L., Bardinelli, C. M., Bowdan, E. S., Cooley, S. S., Crnjar, R. M., Dundulis, E. A., Roitberg, C. A., Spatcher, P. J., Tumlinson, J. H., Weeks, B. L.: Site of production of an oviposition-deterring pheromone components in Rhagoletis pomonella flies. J. Insect Physiol. 28, 1–10 (1982)
Prokopy, R. J., Papaj, D. R., Cooley, S. S., Kallet, C.: On the nature of learning in oviposition site acceptance by apple maggot flies. Anim. Behav. 34, 98–107 (1986)
Roitberg, B. D., Mangel, M.: On the evolutionary ecology of marking pheromones. Evol. Ecol. 2, 289–315 (1988)
Roitberg, B. D., Prokopy, R. J.: Experience required for pheromone recognition by the apple maggot fly. Nature 292, 540–541 (1981)
Roitberg, B. D., Prokopy, R. J.: Host deprivation influence on response of Rhagoletis pomonella to its oviposition deterring pheromone. Physiol. Entomol. 8, 69–72 (1983)
Roitberg, B. D., Prokopy, R. J.: Insects that mark host plants. Bioscience 37, 400–406 (1987)
Rumelhart, D. E., McClelland, J. L., PDP Research Group: Parallel distributed processing. Volume 1: Foundations. Cambridge, MA: MIT Press 1986
Schmidt, J. M., Smith, J. J. B.: Host volume measurement by the parasitoid wasp Trichogramma minutum: The roles of curvature and surface areas. Entomol. Exp. Appl. 39, 213–221 (1985)
Schmidt, J. M., Smith, J. J. B.: Correlations between body angles and substrate curvature in the parasitoid wasp Trichogramma minutum: A possible mechanism of host radius measurement. J. Exp. Biol. 125, 271–285 (1986)
Schmidt, J. M., Smith, J. J. B.: Measurement of host curvature by the parasitoid wasp Trichogramma minutum, and its effect on host examination and progeny allocation. J. Exp. Biol. 129, 151–164 (1987a)
Schmidt, J. M., Smith, J. J. B.: Short interval time measurement by a parasitoid wasp. Science 237, 903–905 (1987b)
Williams, G. C.: Adaptation and natural selection. Princeton, NJ: Princeton University Press 1966
Author information
Authors and Affiliations
Rights and permissions
About this article
Cite this article
Mangel, M. Evolutionary optimization and neural network models of behavior. J. Math. Biol. 28, 237–256 (1990). https://doi.org/10.1007/BF00178775
Received:
Revised:
Issue Date:
DOI: https://doi.org/10.1007/BF00178775