The study of adaptive behavior, including learning, usually centers on the effects of natural selection for individual survival. But because reproduction is evolutionarily more important than survival, sexual selection through mate choice (Darwin, 1871) can also have profound consequences on the evolution of creatures' bodies and behaviors. This article shows, through simulation models, how one type of learning-parental imprinting—can evolve purely through sexual selection, to help in selecting appropriate mates and in tracking changes in the phenotypic makeup of the population across generations. At moderate mutation rates, when population tracking becomes an important but still soluble problem, imprinting proves more useful and evolves more quickly than at low or high mutation rates. We also show that parental imprinting can facilitate the formation of new species. In reviewing the biological literature on imprinting, we note that these results confirm some previous speculations by other researchers concerning the adaptive functions and evolutionary consequences of imprinting. Finally, we discuss how sexual selection through mate choice may have great scientific implications for our understanding of the interactions among evolution, learning, and behavior, and potentially important engineering applications for increasing the efficiency of evolutionary search and optimization methods.
Baldwin, J.M. (1896). A new factor in evolution. American Naturalist, 30, 441-451.
2.
Barlow, G.W., Francis, R.C., & Baumgartner, J.V. (1990). Do the colours of parents, companions and self influence assortative mating in the polychromatic Midas cichlid? Animal Behaviour, 40, 713-722.
3.
Barlow, G.W., & Rogers, W. (1978). Female Midas cichlids' choice of mate in relation to parents' and to own color. Biology of Behavior, 3, 137-145.
4.
Bateson, P. (1978). Sexual imprinting and optimal outbreeding. Nature, 273, 659-660.
5.
Bateson, P. (1983). Optimal outbreeding. In P. Bateson (Ed.), Mate choice. Cambridge , UK: Cambridge University Press.
6.
Bolles, R. C., & Beecher, M. D. (Eds.). (1988). Evolution and learning . Hillsdale, NJ: Erlbaum.
7.
Bowlby, J. (1969). Attachment and loss. New York : Basic Books.
8.
Bradbury, J. W, & Andersson, M. B. (Eds.). (1987). Sexual selection: Testing the alternatives. New York: Wiley
9.
Brandon, R. N., & Burian, R. M. (Eds.). (1984). Genes, organisms, populations: Controversies over the units of selection. Cambridge, MA : MIT Press.
10.
Collins, R.J., & Jefferson, D.R. (1992). The evolution of sexual selection and female choice. In F. J. Varela & P Bourgine (Eds.), Toward a practice of autonomous systems: Proceedings of the First European Conference on Artificial Life. Cambridge, MA: MIT Press/Bradford Books.
11.
Cooch, G. (1961). Ecological aspects of the blue snow goose complex . Auk, 78, 72-89.
12.
Cooke, F., & McNally, C.M. (1975). Mate selection and colour preferences in lesser snow geese. Behaviour, 53, 151-170.
13.
Cosmides, L., & Tooby, J. (1987). From evolution to behavior: Evolutionary psychology as the missing link. In J. Dupre (Ed.), The latest on the best: Essays on evolution and optimality. Cambridge, MA: MIT Press.
14.
Cracraft, J. (1992). Species concepts and speciation analysis. In M. Ereshefsky (Ed.), The units of evolution: Essays on the nature of species. Cambridge, MA: MIT Press/ Bradford Books .
15.
Cronin, H. (1991). The ant and the peacock: Altruism and sexual selection from Darwin to today. Cambridge, UK: Cambridge University Press.
16.
Cushing, J.E., Jr. (1941). Non-genetic mating preference as a factor in evolution. The Condor, 43, 233-236.
17.
Darwin, C. (1871). The descent of man, and selection in relation to sex. London: Murray
18.
Davey, G. (1989). Ecological learning theory. New York: Routledge.
19.
Dawkins, R. (1982). The extended phenotype. Oxford , UK: Oxford University Press.
20.
Dewsbury, D.A. (1981). Effects of novelty on copulatory behavior: The Coolidge effect and related phenomena. Psychological Bulletin , 89, 464-482.
21.
Dobzhansky, T. (1937). Genetics and the origin of species. New York: Columbia University Press.
22.
Eberhard, W.G. (1992). Species isolation, genital mechanics, and the evolution of species-specific genitalia in three species of Macrodactylus beetles (Coleoptera, Scaraceidae, Melolonthinae). Evolution , 46, 1774-1783.
23.
Endler, J.A. (1992). Signals, signal conditions, and the direction of evolution. American Naturalist, 139, S125-S153.
24.
Ereshefsky, M. (Ed.). (1992). The units of evolution: Essays on the nature of species. Cambridge, MA: MIT Press/Bradford Books.
25.
Fisher, H. (1992). Anatomy of love: The natural history of monogamy, adultery, and divorce. New York: Simon & Schuster.
26.
Fisher, R.A. (1930). The genetical theory of natural selection. Oxford, UK: Clarendon Press.
27.
Freud, S. (1950). Totem and taboo ( J. Strachey, Trans., Ed.). New York: W W Nor-ton. (Original work published 1913.)
28.
Goldberg, D.E. (1989). Genetic algorithms in search, optimization, and machine learning. Reading, MA: Addison-Wesley
29.
Gould, J.L., & Gould, C.G. (1989). Sexual selection. New York : Scientific American Library (W.H. Freeman and Co. ).
30.
Grant, B.R., & Grant, P.R. (1979). Darwin's finches: Population variation and sympatric speciation. Proceedings of the National Academy of Sciences of the USA, 76, 2359-2363.
31.
Guilford, T., & Dawkins, M.S. (1991). Receiver psychology and the evolution of animal signals. Animal Behaviour, 42, 1-14.
32.
Harlow, H.F. (1971). Learning to love. San Francisco : Albion.
33.
Harnad, S. (1987). Categorical perception: The groundwork of cognition . New York: Cambridge University Press.
Immelmann, K. (1972). Sexual and other long-term aspects of imprinting in birds and other species. In D. S. Lehrman , R. A. Hinde, & E. Shaw (Eds.), Advances in the study of behavior (Vol. 4). New York: Academic.
39.
Immelmann, K. (1975). The evolutionary significance of early experience . In G. Baerends, C. Beer, & A. Manning (Eds.), Function and evolution in behavior: Essays in honour of Professor Niko Tinbergen . Oxford, UK: Clarendon Press.
40.
Johnston, T.D. (1982). Selective costs and benefits of learning. In J. S. Rosenblatt, R. A. Hinde, C. Beer, & M. C. Busnel (Eds.), Advances in the study of behavior (Vol. 12). New York : Academic.
41.
Kalmus, H., & Maynard Smith, S. (1966). Some evolutionary consequences of pegmatypic mating systems (imprinting). American Naturalist , 100, 619-635.
42.
Kauffman, S. (1993). The origins of order: Self organization and selection in evolution. Oxford, UK: Oxford University Press.
43.
Kirkpatrick, M. (1982). Sexual selection and the evolution of female choice . Evolution, 36, 1-12.
44.
Kirkpatrick, M. (1987). The evolutionary forces acting on female preferences in polygynous animals. In J. W Bradbury & M. B. Andersson (Eds.), Sexual selection: Testing the alternatives. New York: Wiley
45.
Koref Santibanez, S., & Waddington, C.H. (1958). The origin of sexual isolation between different lines within a species. Evolution, 12, 485-493.
46.
Kosswig, K. (1947). Selective mating as a factor for speciation in cichlid fish of East African lakes. Nature, 159, 604-605.
47.
Kosswig, K. (1963). Ways of speciation in fishes. Copeia, 2, 238-244.
48.
Krebs, J. R., & Davies, N. B. (Eds.). (1993). An introduction to behavioral ecology (3rd ed.). Oxford, UK: Blackwell Scientific.
49.
Lande, R. (1981). Models of speciation by sexual selection on polygenic characters. Proceedings of the National Academy of Sciences of the USA, 78, 3721-3725.
50.
Lande, R. (1987). Genetic correlation between the sexes in the evolution of sexual dimorphism and mating preferences. In J. W Bradbury & M. B. Andersson (Eds.), Sexual selection: Testing the alternatives. New York: Wiley.
51.
Littman, M.L., & Ackley, D.H. (1991). Adaptation in constant utility non-stationary environments. In R. K. Belew & L. B. Booker (Eds.), Proceedings of the Fourth International Conference on Genetic Algorithms. San Mateo, CA: Morgan Kaufmann.
52.
Lorenz, K.Z. (1970). Companions as factors in the bird's environment . In K. Z. Lorenz (Ed.), Studies in animal and human behaviour (Vol. 1) (R. Martin, Trans.). Cambridge, MA: Harvard University Press.
53.
(Reprinted from Journal of Ornithology [1935], 83, 137-213)
54.
Marler, P., & Terrace, H. S. (Eds.). (1984). The biology of learning . New York: Springer-Verlag.
55.
Maynard Smith, J. (1978). The evolution of sex. Cambridge , UK: Cambridge University Press.
56.
Mayr, E. (1942). Systematics and the origin of species. New York: Columbia University Press.
57.
Mayr, E. (1947). Ecological factors in speciation. Evolution, 1, 263-288.
58.
McFarland, D. (Ed.) (1987). The Oxford companion to animal behavior. Oxford, UK: Oxford University Press.
59.
Meyer, J.-A., Roitblat, H. L., & Wilson, S. W (Eds.). (1993). From animals to animats 2: Proceedings of the Second International Conference on Simulation of Adaptive Behavior. Cambridge, MA: MIT Press/Bradford Books.
60.
Miller, G.F. (1993a). Evolution of the human brain through runaway sexual selection. Unpublished doctoral thesis, Stanford University Psychology Department, Stanford, CA.
61.
Miller, G.F. (1993b). Artificial life through sexual selection. Artificial Intelligence and Simulation of Behavior Quarterly [special issue on artificial life in the UK]. Manuscript in preparation.
62.
Miller, G.F., & Freyd, J.J. (1993). Dynamic mental representations of animate motion: The interplay among evolutionary, cognitive, and behavioral dynamics. Manuscript submitted for publication.
63.
Miller, G.F., & Todd, P.M. (1990). Exploring adaptive agency I: Theory and methods for simulating the evolution of learning. In D. S. Touretzky, J. L. Elman, T. J. Sejnowski, & G. E. Hinton (Eds.), Proceedings of the 1990 Connectionist Models Summer School. San Mateo, CA: Morgan Kaufmann.
64.
Miller, G.F., & Todd, P.M. (1993a). Evolutionary wanderlust: Sexual selection with directional mate preferences. In J.-A. Meyer , H. L. Roitblat, & S. W Wilson (Eds.), From animals to animats 2: Proceedings of the Second International Conference on Simulation of Adaptive Behavior. Cambridge, MA: MIT Press/Bradford Books.
65.
Miller, G.F., & Todd, P.M. (1993b). The role of mate choice in biocomputation: Sexual selection as a process of search, optimization, and diversification. In Proceedings of the 1992 Monterey Biocomputation Workshop .
66.
Moser, D. (1990). A flight to survival for endangered raptors. Smithsonian, 21, 88-101.
67.
O'Donald, P. (1960). Inbreeding as a result of imprinting. Heredity, 15, 79-85.
68.
O'Donald, P. (1980). Genetic models of sexual selection. Cambridge, UK: Cambridge University Press.
69.
Plotkin, C. G. (Ed.). (1988). The role of behavior in evolution. Cambridge, MA: MIT Press.
70.
Pomiankowski, A., Iwasa, Y., & Nee, S. (1991). The evolution of costly mate preferences. I. Fisher and biased mutation. Evolution, 45(6), 1422-1430.
71.
Pruett-Jones, S. (1992). Independent versus non-independent mate choice: Do females copy each other? American Naturalist, 140, 1000-1009.
72.
Ridley, M. (1987). Imprinting. In D. McFarland (Ed.), The Oxford companion to animal behavior . Oxford, UK: Oxford University Press.
73.
Ryan, M.J. (1990). Sexual selection, sensory systems, and sensory exploitation. Oxford Surveys of Evolutionary Biology, 7, 156-195.
74.
Seiger, M.B. (1967). A computer simulation study of the influence of imprinting on population structure. American Naturalist, 101, 47-57.
75.
Seiger, M.B., & Dixon, R.D. (1970). A computer simulation study of the effects of two behavioral traits on the genetic structure of semi-isolated populations . Evolution, 24, 90-97.
76.
Shepard, R.N. (1987a). Toward a universal law of generalization for psychological science. Science, 237, 1317-1323.
77.
Shepard, R.N. (1987b). Evolution of a mesh between principles of the mind and regularities of the world. In J. Dupre (Ed.), The latest on the best: Essays on evolution and optimality. Cambridge, MA: MIT Press.
78.
Shepard, R.N. (1990). Neural nets for generalization and classification: Comment on Staddon and Reid (1990). Psychological Review, 97(4), 579-580.
79.
Siepen, G., & Crapon de Caprona, M.-D. (1986). The influence of parental color morph on mate choice in the cichlid fish Cichlasoma nigrofasciatum. Ethology, 71, 187-200.
80.
Sluckin, W. (1973). Imprinting and early learning (2nd ed.). Chicago: Aldine.
81.
Spurway, H. (1955). The double relevance of imprinting to taxonomy . British Journal of Animal Behaviour, 3, 123-124.
82.
Staddon, J.E.R. (1983). Adaptive behavior and learning. Cambridge, UK: Cambridge University Press.
83.
Ten Cate, C. (1987). Sexual preferences in zebra finch males raised by two species: II. The internal representation resulting from double imprinting . Animal Behaviour, 35, 321-330.
84.
Thornhill, N.W. (1991). An evolutionary analysis of rules regulating human inbreeding and marriage. Behavioral and Brain Sciences, 14, 247-293.
85.
Thorpe, W.H. (1956). Learning and instinct in animals. Cambridge, MA: Harvard University Press.
86.
Todd, P.M., & Miller, G.F. (1991a). Exploring adaptive agency II: Simulating the evolution of associative learning. In J.-A. Meyer & S. W Wilson (Eds.), From animals to animats: Proceedings of the First International Conference on Simulation of Adaptive Behavior. Cambridge, MA: MIT Press/Bradford Books .
87.
Todd, P.M., & Miller, G.F. (1991b). Exploring adaptive agency III: Simulating the evolution of habituation and sensitization. In H.-P. Schwefel & R. Maenner (Eds.), Proceedings of the First International Conference on Parallel Problem Solving from Nature. Berlin: Springer-Verlag.
88.
Todd, P.M., & Miller, G.F. (1991c). On the sympatric origin of species: Mercurial mating in the Quicksilver model. In R. K. Belew & L. B. Booker (Eds.), Proceedings of the Fourth International Conference on Genetic Algorithms. San Mateo, CA: Morgan Kaufmann.
89.
Trivers, R.L. (1985). Social evolution. Menlo Park, CA: Benjamin/Cummings Publishing.
90.
Vidal, J.M. (1982). 'Auto-imprinting': Effects of prolonged isolation on domestic cocks. Journal of Comparative and Physiological Psychology, 96, 256-267.
91.
Wallace, A.R. (1889). Darwinism: An exposition of the theory of natural selection, with some of its applications. London: Macmillan.
92.
Wiley, E.O. (1992). The evolutionary species concept reconsidered . In M. Ereshefsky (Ed.), The units of evolution: Essays on the nature of species. Cambridge, MA: MIT Press/Bradford Books.
93.
Williams, G.C. (1966). Adaptation and natural selection. Princeton, NJ: Princeton University Press.
94.
Williams, G.C. (1975). Sex and evolution. Princeton, NJ: Princeton University Press.
95.
Wright, S. (1932). The roles of mutation, inbreeding, crossbreeding, and selection in evolution. In Proceedings of the Sixth International Congress on Genetics, 1, 356-366.
