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Psychology Animal Learning
by
Debbie Kelly

Introduction

Understanding why animals, including humans, behave as they do requires an appreciation for learning. Much of the history of animal learning has come from controlled laboratory experiments, coupled with careful field studies of natural behavior. Indeed, there are many approaches to understanding animal behavior, with equally many terms used to describe the endeavor: animal learning, animal cognition, comparative cognition, comparative ethology, and cognitive ethology, to name just a few. However, all these approaches have a foundation built on the principles of learning. Traditionally, psychological approaches to understanding animal learning have been of a general processes nature. This can be seen with the classic Pavlovian, or classical, conditioning experiments of ringing bells and salivating dogs, paradigms known to a broad audience. These principles have been advanced and refined over the years. The field of animal learning has continued to grow and influence other areas of research. The first section of this bibliography provides general Textbooks that cover the topic of animal learning (and cognition), as well as a few specialized textbooks that discuss more-complex forms of cognition. Next, a selection of peer-reviewed Journals that publish in the area of animal learning, cognition, and behavior are presented, with a brief statement of interest. Finally, the remainder provides an introduction to a few central topics of animal learning and ends with a couple of examples of how the foundational understanding of the principles of learning has been applied to the study of specific topics.

Textbooks

The textbooks on animal learning represent an important source for obtaining an overview of topics central to animal learning as well as more-specialized areas of research. Over the years, Domjan has provided an excellent textbook, aimed at undergraduate students, that offers a general overview of topics central to animal learning; now in its sixth edition, it has proven an excellent resource (Domjan 2010). Bouton 2007 is also a textbook directed at an undergraduate and graduate audience. This textbook focuses on the principles of learning, showing how theory and research guide our understanding of these principles. Pearce 2008 gives an introduction to the principles of learning and goes on to examine specific topics of animal cognition—for instance, navigation, imitation, and episodic memory. Papini 2008 is a textbook directed at an undergraduate or graduate audience that takes a different approach from many of the previously discussed textbooks. This textbook integrates important issues in biology and psychology, providing an excellent interdisciplinary resource. Shettleworth 2010 (as well as the previous, 1998 edition) is an excellent textbook that provides a good foundation on animal learning but goes on to examine cognition from the perspectives of comparative cognition, behavioral ecology, and ethology—an excellent synthesis of these areas. Gallistel 1990 is now a classic textbook offering an overview of learning principles, with a focus on time, number, and space. This book not only is accessible to undergraduate students but also is a strong resource for graduate students and researchers alike. Although perhaps now a bit out of date, Mackintosh 1983 is still a very strong resource for those interested in conditioning and associative learning.

  • Bouton, Mark E. 2007. Learning and behavior: A contemporary synthesis. Sunderland, MA: Sinauer.

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    Aimed at midlevel to advanced undergraduate students, this textbook covers the foundational principles of learning and behavior.

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  • Domjan, Michael. 2010. The principles of learning and behavior. 6th ed. Belmont, CA: Wadsworth/Cengage Learning.

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    This textbook, now in its sixth edition, has proven to be a classic in the field. Written for an undergraduate audience, it focuses on the principles of learning and provides an excellent overview of theoretical and empirical research in the area.

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  • Gallistel, C. R. 1990. The organization of learning. Cambridge, MA: MIT Press.

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    This now-classic volume presents an overview of learning principles, with a focus on time, number, and space. Provides excellent coverage of the behavioral and neural basis of learning, with a strong emphasis on spatial learning. Written at the level of advanced undergraduates and graduate students, this book continues to be a valuable resource for those interested in learning.

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  • Mackintosh, N. J. 1983. Conditioning and associative learning. Oxford: Clarendon.

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    An account of the theories and empirical research, focusing on associative learning. This book covers issues within the areas of classical and instrumental conditioning, such as appetitive and aversive reinforcement, laws of association, and excitatory and inhibitory learning, as well as discrimination learning.

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  • Papini, Mauricio R. 2008. Comparative psychology: Evolution and development of behavior. 2d ed. New York: Psychology.

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    This textbook would be an excellent guide for undergraduate students of psychology and biological sciences alike. It is one of the few comparative psychology textbooks that provide a comprehensive introduction to the biological bases of cognition and evolution.

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  • Pearce, John M. 2008. Animal learning and cognition: An introduction. 3d ed. Hove, UK, and New York: Psychology.

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    Written for an undergraduate audience, this textbook gives a basic introduction to the main topics required to understand the principles of learning and behavior. An introduction discusses select topics within the area of animal learning and cognition.

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  • Shettleworth, Sara J. 2010. Cognition, evolution, and behavior. 2d ed. Oxford and New York: Oxford Univ. Press.

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    Excellent interdisciplinary coverage of the field of comparative cognition and learning, easily accessible to students and researchers from a psychology or biology background. This book presents a synthesis of knowledge of psychology, evolution, and behavior in the early 21st century. The material is written in such an integrated and clear manner, this book could be used by senior undergraduate as well as graduate students.

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Journals

There are many peer-reviewed journals that publish articles on animal learning. Many of these venues also include an emphasis on behavior, cognition, and neuroscience. Journals on animal learning range from those focused on the principles, theories, and mechanisms of learning to those that examine animal behavior and cognition. Learning and Behavior and Learning and Motivation publish research articles on the processes involved in learning, behavior, and motivation. Behavioural Processes is a journal that publishes articles across a wide range of topics that report on human and nonhuman behavior. Journal of Experimental Psychology: Animal Behavior Processes is an American Psychological Association (APA) publication that publishes research that examines animal learning and behavior, using experimental methodology. Journal of Comparative Psychology, another APA journal, offers a comparative perspective to the understanding of topics such as learning and behavior. Animal Cognition is a journal that focuses more on cognitive processes and less on learning principles, specifically, but it is an excellent reference for those interested in reading the latest research on the interesting cognitive abilities of animals. Frontiers in Comparative Psychology is another journal with an emphasis on comparative investigations of human, and nonhuman, behavior. Animal Behaviour is a bit distinct from the previously mentioned journals, in that it publishes studies of animal behavior from psychology, behavioral ecology, neuroethology, and biology perspectives. Finally, Neurobiology of Learning and Memory is a good resource for articles that include a neurobiological aspect to the examination of learning and memory.

E-Journals

The presence of e-journals is an exciting new way that one can obtain access to the latest research in the general area of animal learning. Unlike many traditional journals, these two e-journals provide free open access to the published articles, making them available to a wider audience. Comparative Cognition and Behavior Reviews publishes invited reviews on topics related to animal cognition. International Journal of Comparative Psychology publishes research articles exploring topics of human and nonhuman cognition. Some of the articles in this journal include excellent supplemental media in the form of videos or audio clips.

Habituation and Imprinting

Habituation, although not always considered a form of genuine learning, has been a particularly important behavior (and procedure) used in the study of the neurobiology of learning and memory. Thompson and Spencer 1966 is still an excellent resource that clearly outlines the process of habituation. Kandel 2006, which focuses on the author’s Nobel Prize–winning research, not only describes how habituation has been used to study the neurobiology of memory but also gives an excellent context of the history of this research area. Imprinting was once thought to be a special form of learning that was instantaneous and irreversible; the classic example is the goslings of Konrad Lorenz. However, these assumptions are being challenged; a good review of this literature is provided in Bolhuis 1991. Similar to habituation, the study of imprinting has led to a greater understanding of the neural basis for learning, as reviewed in a comprehensive way in Horn 2004.

  • Bolhuis, Johan J. 1991. Mechanisms of avian imprinting: A review. Biological Reviews of the Cambridge Philosophical Society 66.4: 303–345.

    DOI: 10.1111/j.1469-185X.1991.tb01145.xSave Citation »Export Citation »E-mail Citation »

    A comprehensive review of the literature on avian imprinting. Bolhuis discusses studies that suggest imprinting is a form of associative learning. Also provides evidence that sexual and filial imprinting is reversible, although preferences for earlier imprinted stimuli are stronger than for later stimuli.

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  • Horn, Gabriel. 2004. Pathways of the past: The imprint of memory. Nature Reviews Neuroscience 5:108–120.

    DOI: 10.1038/nrn1324Save Citation »Export Citation »E-mail Citation »

    A comprehensive review of the literature on the neural underpinnings of imprinting. Studies examining the changes in the intermediate and medial hyperstriatum ventrale are presented and discussed.

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  • Kandel, Eric R. 2006. In search of memory: The emergence of a new science of mind. New York: Norton.

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    A popular science book that provides an excellent, easy-to-read history of Kandel’s Nobel Prize–winning research on memory. Presents several chapters on neural bases of habituation, using the classic model, Aplysia.

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  • Thompson, R. F., and W. A. Spencer. 1966. Habituation: A model phenomenon for the study of neuronal substrates of behavior. Psychological Review 73.1: 16–43.

    DOI: 10.1037/h0022681Save Citation »Export Citation »E-mail Citation »

    A “guidebook” to the nine properties of habituation.

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Associative Learning

Associative learning is a process by which an animal can learn about the predictive nature of events within an environment. Associative learning can be measured when an animal’s behavior changes as a result of event pairings. Two methods for examining associative learning are classical, or Pavlovian, conditioning and instrumental conditioning. Mackintosh 1983 presents a succinct overview of the important theories of classical and instrumental conditioning. Fanselow and Poulous 2005 offers a careful review of two classical procedures used in associative learning and explains the underlying neural basis of these paradigms. Gallistel 1990 is now a classic text covering the processes of learning and the formation of representations for time, number, and space.

  • Fanselow, Michael S., and Andrew M. Poulos. 2005. The neuroscience of mammalian associative learning. Annual Review of Psychology 56:207–234.

    DOI: 10.1146/annurev.psych.56.091103.070213Save Citation »Export Citation »E-mail Citation »

    This review provides a discussion of the two classic techniques for studying associative learning in mammals, fear and eyeblink, as well as the brain circuits involved in these conditioning paradigms.

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  • Gallistel, C. R. 1990. The organization of learning. Cambridge, MA: MIT Press.

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    This now-classic volume presents an overview of learning principles, with a focus on time, number, and space.

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  • Mackintosh, N. J. 1983. Conditioning and associative learning. Oxford: Clarendon.

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    Mackintosh provides a detailed examination of the theoretical approaches and empirical evidence of associative learning. Several topics are covered, such as appetitive and aversive reinforcement, the laws of association, and excitatory and inhibitory learning. This book remains a “must read” for those interested in classical and instrumental conditioning.

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Classical, or Pavlovian, Conditioning

Pavlovian conditioning is one means of investigating how animals learn about the predictability of events. During Pavlovian conditioning, at least two events are paired together: a neutral conditioned stimulus (CS) and a biologically relevant unconditioned stimulus (US). The close temporal pairing of a CS (e.g., a light) with the delivery of a US (e.g., food) leads to the animal emitting an unconditioned response (UR; e.g., salivation); with repeated pairings, the CS itself comes to elicit responses that are now termed conditioned responses (CR)—even when the US is not presented. Rescorla 1988 provides a clear description of how the field of associative learning has changed and grown since its initial studies. A more current analysis provided in Miller 2006 argues that the field of associative learning still has many issues to face and presents three main challenges to the associative models.

Instrumental, or Operant, Conditioning

Instrumental conditioning is another means of investigating how animals learn about the predictability of events. An animal can learn about events through the presentation of rewards and punishments that result from the performance of a behavior, with behavior being repeated following a reward and decreased following a punishment. The original research, classic studies of instrumental conditioning still worth reading about, is from Thorndike (Thorndike 1898), who gives simple to fairly elaborate puzzle boxes to animals and describes their ability to solve these puzzles. One interesting phenomenon is the transfer from Pavlovian conditioning to instrumental conditioning; this is discussed in Holmes, et al. 2010 from a neurobehavioral perspective.

  • Holmes, Nathan M., Alain R. Marchand, and Etienne Coutureau. 2010. Pavlovian to instrumental transfer: A neurobehavioural perspective. Neuroscience and Biobehavioral Reviews 34.8: 1277–1295.

    DOI: 10.1016/j.neubiorev.2010.03.007Save Citation »Export Citation »E-mail Citation »

    This article provides a description of the theories of Pavlovian-to-instrumental transfer (PIT) and the underlying neural mechanisms and presents empirical evidence that the order of training is an important variable.

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  • Thorndike, Edward L. 1898. Animal intelligence: An experimental study of the associative processes in animals. Psychological Review, Monograph Supplements 2.4. New York and London: Macmillan.

    DOI: 10.1037/10780-000Save Citation »Export Citation »E-mail Citation »

    This article is a classic read for anyone interested in the investigation of animal intelligence and learning. It provides an in-depth report, by Thorndike, of the learning strategies of animal subjects as they attempt to solve puzzle boxes designed to range in difficulty.

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Discrimination Learning

The ability to distinguish between two or more items is important to many animals. Such discriminations may be between items such as edible and nonedible foods or conspecifics and predators. Discriminative behavior can be seen along a continuum from that which does not require training to highly complex forms of category discrimination. In a classic study of sign stimuli in animals, Hailman 1967 showed that herring gull chicks, which peck at a red spot on their parent’s bill to stimulate regurgitation, will readily peck at an artificial red stimulus, such as a knitting needle. However, as the chicks mature, their behavior becomes more discriminating and is influenced by additional, more subtle, cues, such as the shape of the head area. A complex form of discriminative behavior shown most often by pigeons and primates is that of same/different category learning. An excellent discussion of whether pigeons (and other nonhuman animals) are categorizing these items or using perceptual variability to solve the problem (or indeed, impressive feats of memorization, as shown in Cook and Fagot 2009) is presented in the volume Cook 2001 as well as a collection of articles in Wasserman and Zentall 2006.

  • Cook, Robert G. 2001. Avian Visual Cognition. Boston: Comparative Cognition.

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    This cyberbook presents a series of reviews from prominent researchers in the area of animal discrimination learning and classification.

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    • Cook, Robert, and Joel Fagot. 2009. First trial rewards promote 1-trial learning and prolonged memory in pigeons and baboons. Proceedings of the National Academy of Sciences of the United States of America 106.23: 9530–9533.

      DOI: 10.1073/pnas.0903378106Save Citation »Export Citation »E-mail Citation »

      Pigeons and baboons were able to learn hundreds of pictures after a single picture-reinforcement exposure; these learned acquisitions were present for several months after initial learning.

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    • Hailman, Jack P. 1967. The ontogeny of an instinct: The pecking response in chicks of the laughing gull (Larus atricilla L.) and related species. Leiden: E. J. Brill.

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      A now-classic work that investigates the behavioral responses of chicks to sign stimuli; this research shows that initially, newly hatched chicks’ responding is focused on the red color spot on the lower mandible of the parent’s bill, and this response behavior can be elicited by quite artificial stimuli. However, with maturation the chicks are influenced by more of the stimulus, such as the shape of the parent’s head, and thus show an increase in responding to more-naturalistic gull models.

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    • Wasserman, Edward A., and Thomas R. Zentall. 2006. Comparative cognition: Experimental explorations of animal intelligence. Oxford and New York: Oxford Univ. Press.

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      This comprehensive edited book presents a series of reviews from prominent researchers in the area of animal cognition. Chapters 1, 2, and 6 are particularly informative for the understanding of discrimination learning and categorization by animals.

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    Spatial Learning

    Many animals must be able to locomote from one position to another, in the search of nourishment, mates, and home sites, for example. The study of spatial learning examines what types of cues an animal can use when needing to relocate from one position to the next. Perhaps not surprisingly, animals use many different cues to orient and navigate within an environment, and these cues depend not only on the species, but also the distance to be traveled and the amount and nature of spatial cues available. The complexity of cue use ranges from dead reckoning (or path integration) to the integration of multiple landmarks and geometry use. An excellent collection of articles on spatial learning is presented in Healy 1998 as well as the e-book Brown and Cook 2006. A now-classic examination of the role of the hippocampus in spatial learning is explored in O’Keefe and Nadel 1978. This publication has generated considerable research in the area of spatial learning, because the authors propose that there is a special spatial learning system that is distinct from associative learning. The idea that geometry may be modular within spatial learning has also generated an incredible amount of research since the mid-1980s. Although there have been studies with results calling into question the geometric module, Cheng’s work in this area (Cheng 1986) has been pioneering.

    • Brown, Michael F., and Robert G. Cook. 2006. Animal Spatial Cognition: Comparative, Neural, and Computational Approaches. Boston: Comparative Cognition.

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      A wide, sweeping collection of reviews on spatial learning and cognition. Provides an overview of topics such as approaches to the study of spatial cognition, geometric learning, neural underpinnings of spatial representations, and human spatial learning, to name a few. Excellent illustrations accompany many of these chapters.

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      • Cheng, K. 1986. A purely geometric module in the rat’s spatial representation. Cognition 23.2: 149–178.

        DOI: 10.1016/0010-0277(86)90041-7Save Citation »Export Citation »E-mail Citation »

        Pioneering research showing that rats strongly rely on the geometric properties of an environment when searching for a hidden goal; the rats show a primary encoding of geometry even when distinctive features are available and provide a more accurate cue. This article has generated a plethora of research into the idea of a geometric module and the encoding of geometry across a wide range of species.

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      • Healy, Sue, ed. 1998. Spatial representations in animals. Oxford and New York: Oxford Univ. Press.

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        A selection of articles focused on providing information as to how animals orient and navigate within their environments. The majority of these articles examine spatial learning at a behavioral level, although the final chapter presents a nice overview of the neural bases of spatial representations.

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      • O’Keefe, John, and Lynn Nadel. 1978. The hippocampus as a cognitive map. Oxford: Clarendon.

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        This classic work brings together research investigating the function of the hippocampus and its core role as a memory system that provides a spatial “map” of an animal’s environment.

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      LAST MODIFIED: 11/29/2011

      DOI: 10.1093/OBO/9780199828340-0073

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