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LEARNING AND MOTIVATION 15, 321-333 (1984)

An Ecological Approach to Learning

WILLIAM TIMBERLAKE

Indiana University

This paper introduces a special journal issue devoted to the study of learning in ecological and developmental contexts. In the past, the dominant approach to the study of learning has depended upon the choice of arbitrary problems, stimuli, and responses to guarantee the generality of principles discovered in the laboratory. Recently, though, there has been considerable interest in integrating an ecological (functional) account of learning with the strengths of the classic laboratory approach. One form of integration is to use the technology of the laboratory to look for the operation of general learning laws in ecological problems. The approach favored here is to treat learning as a biological phenomenon by first placing it within a functional system of behavior, and then by analyzing where and how learning modifies the operation of that system. Because the results of such analyses are defined with respect to functioning systems rather than procedural paradigms, the ecological approach readily makes contact with issues in evolution, development, and physiology, an ability not completely shared by the classic general-process approach. Concerns about laboratory versus field, function versus mechanism, generality of results, and adaptive “storytelling” can be resolved or further clarified by the present approach. The papers in this issue represent a cross-section of research stemming from an ecological approach to learning, and provide specific analyses of how learning modifies and is expressed in functional systems of behavior. 0 1984 Academic Press. Inc.

At the turn of the century the study of animal learning was divided into two camps: the arbitrary general-process approach, those who believed that learning was best studied through the use of artificial problems involving arbitrarily chosen (generic) stimuli and behaviors; and the ecological approach, those who argued that learning should be studied in the context of natural problems, stimuli, and behavior. (The interested reader is referred to Galef, 1984, and Timberlake, 1983b, for further consideration of this conflict.) The arbitrary approach rapidly assumed

Preparation of this manuscript was supported in part by NSF Grant 82-10139 and PHS Grants MH 37892 and MH 39345. I thank Jeffrey Alberts, Don Gawley, David Gubemick, Eliot Hearst, Karen Hollis, and especially Gary Lucas for comments, and Michael Rashotte for referring me to Watson. Requests for reprints should be sent to Dr. William Timberlake, Department of Psychology, Indiana University, Bloomington. IN 47405.

321 0023-9690/84 $3.00

Copyright 0 1984 by Academic Press. Inc. All rights of reproduction in any form reserved.

322 WILLIAM TIMBERLAKE

dominance because (1) it clearly separated the study of learning from increasingly questionable explanations of behavior in terms of the animal’s mental life or instincts, (2) it focused on the development of general laws and theories presumably applicable to all stimuli and responses, and (3) it encouraged the development of standardized tasks and paradigms that were thought to allow comparison of the abilities of different species and extrapolation to humans.

After a long period of successful development and expansion to other areas and disciplines, the arbitrary general-process approach began to lose ground in the 1950s. At a theoretical level, increasingly complex and indefinite concepts were needed to account for existing data. At an empirical level, researchers began to note important differences in the way learning manipulations affected different responses and species (e.g., Bolles, 1970; Breland & Breland, 1961, 1966; Garcia & Koelling, 1966; Garcia, McGowan, & Green, 1972; Rozin & Kalat, 1971; Seligman, 1970; Shettleworth, 1972).

The ecological concerns summarily dismissed at the turn of the century gradually have reappeared and now threaten to divide the field again. The arbitrary general-process approach has on its side the weight of tradition and cumulated data seemingly confirming the sufficiency of general laws to account for all learning (e.g., Mackintosh, 1974). Apparent contradictions of these laws have been accommodated within the category of quantitative constraints on their operation (e.g., Domjan, 1983; Logue, 1979). On the other side, the ecological approach and its emphasis on species- and problem-specific variation in learning have powerful allies in newly developed fields such as behavioral ecology, sociobiology, and a revitalized area of animal development. The ecological approach also has profited from increased field data (Galef, 1984), and a greater so- phistication concerning the role of genetics and evolution (e.g., Johnston, 1981, 1982; Kovach, 1984; Plotkin & Odling-Smee, 1979). Given the apparent intransigence of the two sides, there appears to be the possibility of a permanent split in the study of learning similar to that now threatening learning and development (e.g., Gottlieb, 1983; Miller & Blaich, 1984).

However, along with the obvious conflict there has emerged a significant interest in finding a common ground between the arbitrary and ecological approaches. Some of this interest has come from researchers influenced by the biological sciences who have begun to appreciate the potential role of learning in fitness, evolution, and development (e.g., Alberts & Gubernick, 1984; Baker, Tomback, Thompson, Theimer, & Bradley, 1984; Emlen, 1973; Hollis, Martin, Cadieux, & Colbert, 1984; King & West, 1984; Kroodsma, 1982; Marler & Peters, 1982; Mayr, 1974; Plotkin & Odling-Smee, 1979). Interest in a common ground has also been expressed within the arbitrary tradition, as researchers have struggled to deal with the species-typical results produced by the supposedly arbitrary

ECOLOGICAL APPROACH TO LEARNING 323

problems and stimuli of the laboratory: autoshaping, polydipsia, misbehavior, taste-aversion learning, and systematic constraints on learning and performance (e.g., Bolles, 1970; Domjan, 1980, 1983; Falk, 1971; Hollis, 1982; Logue, 1979; Seligman, 1970; Shettleworth, 1972, 1983; Staddon, 1983).

Though the common interest in integration within the ecological and arbitrary general-process approaches speaks well for the possibility of cooperation, the actual combination of the approaches has been slow to develop. Researchers in the arbitrary general-process tradition have been uncertain as to how to deal with nonstandard paradigms that use natural stimuli and responses in studying ecological problems. There is still a fear that effort on such problems will be wasted because it will consist merely of endless “botanizing” of responses, stimuli, and reinforcers with no general integrative theory (Skinner, 1938). On the other hand, researchers in the more biological tradition find it difficult to fit the learning problems they study into the paradigms and procedures of the arbitrary approach. For many it is difficult to understand why one would study abstract principles that have no ecological or evolutionary grounding.

THE APPLICATION OF LEARNING TECHNOLOGY TO ECOLOGICAL PROBLEMS

The most frequent technique used to bridge the gulf between ecological considerations and laboratory research has been to apply the principles and techniques of the laboratory to the analysis of particular ecological problems. The most common procedure has been to examine simulations of real-world problems for the possible contribution of the principles of Pavlovian or operant conditioning established in the laboratory. This procedure has spawned explanations of and research on several real- world problems, ranging from imprinting (Hoffman & Segal, 1983) through foraging (e.g., Collier, 1983; Collier & Rovee-Collier, 1981; Fantino & Abarca, in press) to economic behavior (e.g., Allison, 1983; Rachlin, Battalio, Kagel, & Green, 1981; Rachlin, Green, Kagel, & Battalio, 1976).

A related procedure has been to use the procedures and equipment developed in the laboratory to analyze the determinants of a particular type of ecological learning. At several recent interdisciplinary conferences, it has been suggested that if biologists will clearly state what they want to know, psychologists will design the apparatus and experiments to find out the answers. Though there is obviously more to an integrative approach to learning than this, the statistical, methodological, and conceptual facility developed within the laboratory tradition provides powerful tools for the analysis of mechanism (and function). The contribution of these tools can be seen especially clearly in work on foraging (e.g., Kamil & Yoerg, 1982; Pietrewicz & Kamil, 1981; Olton, Handelman, & Walker, 1981).

Among some researchers, though, there remains a feeling that biologists and psychologists still are talking past each other. The application of


laboratory techniques to species-typical problems does not in itself provide a general ecological framework for the study of learning. Psychologists remain preoccupied with a general-process view of learning mechanisms independent of a particular functional context. As a result, major aspects of phenomena such as bird song learning, imprinting, the development of feeding behavior, and even classic subjects such as the nature of mind in apes (e.g., Premack, 1976), have remained peripheral to the laboratory study of learning. When confronted with real gaps in our knowledge, proponents of the arbitrary tradition fall back on the argument that examples of learning in nearly all species can be explained within current paradigms and theories of learning, and that all other learning phenomena can in principle be explained using the same approach. They attribute the lack of progress in dealing with particular examples of ecological learning to the complexity of the situation, the absence of good field data, and the limited possibilities of control of the circumstances.

AN ECOLOGICAL (FUNCTIONAL) APPROACH TO THE STUDY OF LEARNING

A more general solution is to take learning as a biological phenomenon to be analyzed and explained in the same way that territorial or feeding behavior is analyzed and explained (Rozin & Kalat, 1971). An ecological approach as identified here seeks simply to set the study of learning within the functional context in which it evolved. Instead of presuming that learning de novo produces the basic behavior of the organism, the ecological approach begins with the assumption that the organism comes equipped with organized stimulus sensitivities, processing proclivities, response structures, and integrative states evolved to produce adaptive behavior in particular environments (e.g., Timberlake, 1983a. 1983b). Learning occurs as modifications in the operation, inclusion, and linkage of different mechanisms within such a functional context (cf. Nottebohm, 1972).

It is important to emphasize that learning is not the source of a functional system. That is, learning does not provide the fundamental structure or the types of mechanism, or determine the basis of its own opportunities for occurrence. From the standpoint of evolution and development, a functional system precedes, underlies, and determines much of the nature of learning. As William James (1890) noted, instincts both precede and provide the opportunity for learning, or, in the words of the Breland and Breland (1966), “To view animal behavior within a framework of learning is to miss a great deal of nature’s basic program or format.” An ecological analysis of learning considers how, where, and to what end the operation of a basic functional system is modified by experience.

Most importantly, placing learning within a functional context consid- erably improves the possibility of integrating the study of learning with the study of development, physiology, and genetics (e.g., Alberts &


Gubernick, 1984; Davis, 1984; Kovach, 1984). Learning is no longer viewed as a single unique connection or representation located in some so far undiscovered part, pathway, or process of the brain, or a concept located even more mysteriously in the experimenter’s procedures. Instead, learning is a modification in the operation of a functional system at the level of stimulus control, processing tendencies, motor patterns, or integration and regulation. Learning is not limited to particular a priori forms, but is constrained within functional systems and by problem- specific mechanisms (e.g., Alberts & Gubemick, 1984; Davis, 1984; Hogan, 1984). Moreover, it is not necessary to force learning into conceptual- procedural categories, such as instrumental and Pavlovian conditioning, in order to analyze its determinants.

It is my belief, and one which I think is shared by the contributors to this issue, that a great deal of headway can be made by careful analyses of where and in what fashion learning modifies the operation of the functional systems that support adaptive behavior. The results of such studies should be more easily related to issues of ontogeny, physiology, and evolution than the results of traditional laboratory analyses of learning. Equally important, in the long run the development of a general theory of learning actually will be advanced rather than inhibited by such careful botanizing.

Do these arguments mean we should abandon the great store of in- formation contained in the traditional laboratory study of learning? Certainly not to the extent that the techniques, skills, and concepts developed in laboratory work are useful in analyzing the mechanisms of particular functional systems. Perhaps not at all, for as I have argued elsewhere (Timberlake, 1983a, 1983b), the allegedly arbitrary analysis of animal learning has been arbitrary primarily at the level of vocabulary and concepts. At the level of procedures and apparatus design, good experimenters have been highly successful in creating at least semiappropriate ecological contexts for the study of learning.

The majority of laboratory research can be viewed as investigations of learning within the functional context of the feeding and avoidance systems of a few species. Most often by trial and error, experimenters have arranged the experimental environment to allow them to present stimuli and measure responses relevant to these systems, but with in- complete awareness or acknowledgment of what has been done. It seems most appropriate that we now recognize this experimental art and the artifice that hides it. Previous laboratory work contains considerable knowledge about learning in functional systems that is embedded in particular combinations of apparatus, procedures, and results. In many cases it may be possible to decipher the mechanisms that have actually been under study (e.g., Timberlake, 1983a; Timberlake B Lucas, 1984).


SOME BOTHERSOME ISSUES

Field versus Laboratory and Function versus Mechanism

The distinction between field and laboratory approaches is not identical to that between the study of function and mechanism. However, in general the split in emphasis has traditionally been along the same lines in each of these distinctions. The question of importance at this point is whether the conflicts represented by these distinctions can be resolved within the present ecological approach.

In a general way the solution to the conflict between laboratory and field research has been clear at least since Watson wrote in his Introduction to Comparative Psychology. “It would seem obvious there is no conflict between field work and laboratory work. The field is both the source of problems and the place where the laboratory solutions of these problems are tested” (Watson, 1914, p. 31). Despite this proffered resolution, feelings still run high on the perceived relative merits of the two approaches. It is as though each new generation of investigators must debate the issue again, possibly because of the lack of a lasting conceptual integration of the two, and possibly because of something as fundamental as personality differences in the investigators (Beer, 1980).

From the present point of view there is no necessary conflict between laboratory and field research or between the study of mechanism and function (Shettleworth. 1983). Since the focus of an ecological approach is on phenotype and survival, the field and function are obviously important referents. However, adaptive function results only from the operation of particular mechanisms, and frequently the best way to study mechanisms is under laboratory conditions. A key to rapid progress in research is to construct a controlled set of circumstances in which a functional system operates in an ecologically understandable way. Whether this set of circumstances occurs inside a box, or outside in the sunshine and rain, is not the critical issue. What is critical is that the test environment and procedures elicit and support the critical components of the system under study. If the situation is too barren or too complex, learning may not be expressed in typical or even recognizable form. Viewed in this way, there is no dichotomy between laboratory and field or between function and mechanism. Each represents a reasonable concern in studying a particular functional system.

There are, though, some important issues that inevitably bedevil glib attempts to integrate laboratory and field research and the study of mechanism and function. First, for several reasons the mechanisms one studies in the laboratory may have little to do with the ecological function of learning. Because of the control possibilities and power of the laboratory, researchers are easily shaped by their results to focus on a particular relation of procedures and outcomes. Yet these relations may not reflect


the operation of learning mechanisms in ecological circumstances. Thus, because laboratory research often analyzes capacity rather than typical operation, reliable determinants of behavior in the laboratory may account for little of the behavioral variance in the animal’s selection environment (see also Galef, 1984).

Thus, if we viewed the results of controlled laboratory-type research in isolation, we might conclude that male sticklebacks court females four times their own size, young gulls commonly peck at knitting needles, adult gulls sit on football-sized eggs, and rats are miniature economists with beady eyes and fur suits. Only when we consider the ecological context is it possible to begin to understand the selection of the mechanisms and their milieu of operation. Watson (1914) addressed this issue by noting that after years of studying the senses of birds in a laboratory, one would predict with confidence the utter absurdity of a bird’s finding its way home when carried out to sea a great distance. He continued, “the incompleteness of the laboratory study would be discovered only when we, so to speak, began to put the bird together again” (Watson, 1914, p. 31).

There are also dangers, though, in elevating field results beyond what actually has been established. In the field there is an almost overwhelming tendency to view outcomes and stimuli through the eyes of the experimenter rather than through the mechanisms that produce the processing and behavior. The adaptiveness of behavior in the field does not require the level of decision mechanism frequently assumed, and almost certainly does not entail the sorts of optimality and fitness calculations currently popular in explanations of field data. As pointed out by Watson (1914), there is no guaranteed technique for inferring mechanism from function, and as pointed out by Galef (1984) there is not even the necessity that a mechanism producing a particular functional outcome in the laboratory is the same one producing that outcome in the field. A particular environment or the motivational state or history of the organism is often a significant mediator of the connection between function and mechanism. Sometimes a modest amount of laboratory research can suggest the unreasonable nature of critical assumptions about the basis of naturally occurring behavior (e.g., Timberlake, 1984).

In general, the techniques of laboratory and field and the focus on mechanism or function are simply different and necessarily dependent ways to get at the nature of learning and behavior. What drives the best research is the question, not the procedures or beliefs of the scientist. The power of the laboratory is most useful when mechanisms have been identified that are relevant to a known and measurable function. Then one can take control of the stimulus and organismic conditions to analyze the mechanisms and precisely measure their function (e.g., Baerends & Drent, 1982; Griffin, 1958).


The Generality Problem

An apparently common fear of researchers in learning, possibly stemming in subterranean fashion from our perceived lack of influence on the world or even on each other. is that they will waste their research time developing explanations of phenomena so esoteric that few will care. This fear seems to hold researchers transfixed at the level of studying and interpreting particular examples of learning only within the framework of an arbitrary general-process approach to learning.

As far as I can see this approach is guaranteed to produce a high proportion of isolated facts and theories because the situation is conceived inappropriately in generic rather than species-specific terms. A basic, universal learning mechanism to which we are so wont to refer may be “in there” in some form (e.g., Hawkins & Kandel, 1984), but evolution selects directly for outcomes using the entire functional set of mechanisms on hand, rather than for some single, basic associative process (e.g., Davis, 1984). Learning is embedded by selection in an already functioning context, not tacked on like an extra appendage suited for an entirely new task.

There is only one certain general law of learning. Learning evolved as selected modifications of mechanisms already serving a particular function, and it occurs where the typical result of such modifications has enhanced the reproductive fitness of the organism. Any other law ultimately must be compatible with this law. The number of learning processes and mechanisms that exist and their generality is a matter of empirical research. If a general law of association exists across all phyla, it implies the existence of a common selection pressure; it most likely does not imply the existence of a common mechanism.

The simplest interpretation of the experiments that support general learning processes is that experimenters have focused on procedures, problems, and measures that make contact with similar ecological functions and thus produce robust and reliable data (e.g., Timberlake, 1983a, 1983b). In less successful experiments, researchers have not made good contact with aspects of the animals ecology that fit their models. Thus, they have encountered either weak phenomena, or robust but unexplained phenomena, such as misbehavior (Breland & Breland, 1961; Timberlake. Wahl, & King, 1982), superstition (Skinner, 1948; Staddon & Simmelhag, 1971; Timberlake & Lucas, 1984) or adjunctive behavior (Falk. 1971).

I think it quite likely that the differences between Pavlovian and operant conditioning, issues of exposure learning versus reinforcement, and anomalous phenomena such as partial reinforcement will be clarified and integrated in a larger theory once it is adequately recognized that we are studying a functioning organism rather than an isolated general learning mehanism. Responses, motivational disequilibriums, and stimulus sensitivities are not setting conditions for learning, they are all parts of


an integrated “package” that was selected as an operating unit. Some learning effects appear to reflect the operation of a general process, such as discrimination or generalization. However, a more careful consideration reveals the importance of the complete functional system. For example, only a brief stimulus exposure at an early age in a duckling will produce extensive effects on later reproductive behavior (Hess, 1973). Evolution does not select for a storage mechanism alone, or for any association, discrimination, or generalization process in isolation from behavior and ecological context. Evolution always selects for outcomes in particular organismic and environmental circumstances.

The Problem of Adaptive Storytelling

An often noted flaw of functional accounts of behavior is the tendency to tell face-valid stories about the presumed adaptive nature of a particular organismic feature in the absence of any attempt to verify the basis of the story (Bitterman, 1975; Gould, 1980). The present attempt to study learning within an evolved functional context potentially encounters similar problems. Without denying the dangers of such storytelling, several points seem worth making. First, it should be noted that storytelling is not the exclusive province of functional accounts. Accounts in terms of mechanisms are also subject to the dangers of unbridled speculation, for example, the invoking of hunger centers in the brain as an explanation for feeding, or the unchecked use of reinforcement as an explanation for behavior change. As I have noted elsewhere:

The widespread use of reinforcement concepts to account for behavior developed as an alternative to careless explanations couched in terms of instincts. The abandonment of instinct as an explanatory device was easy because the concept of instinct was largely an unanalyzed assumption. However. now may be an appropriate time to look more carefully at the careless use of reward to explain behavior. In a sense we have simply switched from explaining behavior in terms of inadequately analyzed causal constructs located in the organism to inadequately analyzed causal constructs located in the experimenter’s procedures. (Timberlake. 1983a, p. 214)

Second, “stories” about phenomena are basically a first step in developing theories. If a nascent theory cannot generate a story that can be applied to a broader range of phenomena, it probably is not worth pursuing. At the same time, stories are just an initial step. The key to moving beyond stories is the interlocking analysis of function and mechanism. This is precisely the type of research encouraged by an ecological approach to the study of learning.

THE PRESENT PAPERS

The present papers share a commitment to analyzing learning as an evolved phenomenon. The majority focus on particular mechanisms of


expression within a functional context. Alberts and Gubernick, Hogan, and Davis are concerned with learning in the functional context of feeding in rat pups, chicks, and carnivorous marine molluscs, respectively: Kovach, and Miller and Blaich deal with perceptual learning in the context of parent-young interaction in quail and ducks; Baker et al., and King and West explore the potential role of song learning in reproduction and speciation in white-crowned sparrows and cowbirds; Hollis et ul. are concerned with the contribution of excitatory and inhibitory learning to territory defense in anabantid fish; Galef is concerned primarily with clarifying the complex interrelations of laboratory and field work.

Except for a few areas, these papers form a reasonable cross section of the sort of research being conducted today in a functional ecological approach to learning. The area of foraging has been slighted unintentionally because invited contributors had previous commitments that prevented their participation. A disproportionate number of the papers (seven of nine) are related to ontogeny, but I find this no accident. The study of development demands that one recognize explicitly the preexisting structures of the organism and the functional context in which learning occurs. Interestingly, only one contributor publishes predominantly in animal learning journals, but all clearly have much to offer specialists in terms of data, paradigms, and questions. I would like to express my thanks to these authors for their willingness to stick to a timetable in order to share their research with a wider audience. I think their contribution to this audience has been considerable, and I hope others find it likewise.

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Received October 18, 1984

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An Ecological Approach to Learningbsl/Papers from lab/Timberla_1984d.pdf · An Ecological Approach to Learning ... A related procedure has been to use the procedures and equipment