Syllabus Psychology e-1090: Animal Cognition




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  1. Syllabus Psychology E-1090: Animal Cognition

This syllabus is tentative—depending on what papers are published during the course, lectures may be altered to include newer material. Also, given class interests, some topics may be given more or less time than indicated. Some lectures will concentrate on a particular species, other lectures will concentrate on a topic.


Grades will be partially dependent upon class interaction. Depending on class size, I may or may not get to know each student individually, but there will be at least 20-30 mins in each class set aside for Q&A and comments, and students are expected to have read the assigned material and be prepared for some discussion after the lecture. Those who regularly participate in a positive manner (i.e., not just to hear themselves speak) will likely find some ‘extra credit’ on their grades.
Grades will mostly depend on take-home midterm and final essays. The midterm will be an assigned reading of a published manuscript that students will be required to critique with respect to sound experimental design and interpretation of the data. The class will vote on whether the midterm is due before or after spring break. The final will be an exercise in designing an original experiment to determine a cognitive capacity of some animal species.
The topic of animal cognition covers an immense range of topics and species. We will do little more than scratch the surface…but I hope that the representative topics and papers will at least serve to excite interest in the field.

Meeting 1: Introduction to the topic. A lecture on “What is cognition?” No assigned reading. A brief ‘writing assignment’ is required by the Extension School.

Meeting 2: Categorization: The lab versus nature. Clearly animals categorize their world… prey and predator, mate and nonmate, kin and not-kin, edible and nonedible. To what extent are these categories learned? Do they reflect cognitive processes? We will examine various species and various types of categorization.
Khallad, Y. Conceptualization in the pigeon: What do we know? International Journal of Psychology 39, 2004, 73-94.

Pepperberg, I.M. Cognition in the African Grey parrot: Preliminary evidence for auditory/vocal comprehension of the class concept. Animal Learning & Behavior 11,1983, 179-185.

Bovet, D., Vauclair, J. Functional categorization of objects and of their pictures in baboons (Papio anubis). Learning and Motivation 29, 1998, 309-322. [Note correction, vol. 30, p. 128]

Dépy, D., Fagot, J., Vauclair, J. Categorisation of three-dimensional stimuli by humans and baboons: Search for prototype effects. Behavioural Processes 39,1997, 299-306.



Meeting 3: Concepts of same-different. An abstract level of categorization. Why some studies purporting to have shown same-different probably have not, and why the topic is important.
Premack, D. On the abstractness of human concepts: Why it would be difficult to talk to a pigeon. In Cognitive Processes in Animal Behavior, SH Hulse, H Fowler, WK Honig, Eds., pp. 421-451. Hillsdale, NJ: Erlbaum 1978.

Pepperberg, I.M. Acquisition of the same/different concept by an African Grey parrot (Psittacus erithacus): Learning with respect to categories of color, shape, and material. Animal Learning & Behavior 15, 1987, 423-432.

Cook, R.G., Cavoto, B.R., Katz, J.S., Cavoto, K.K. Pigeon perception and discrimination of rapidly changing texture stimuli. Journal of Experimental Psychology: Animal Behavior Processes 23,1997, 390-400.

Cook, R.G., Katz, J.S., Cavoto, B.R. Pigeon same-different concept learning with multiple stimulus classes. Journal of Experimental Psychology: Animal Behavior Processes 23, 1997, 417-433.

Wasserman, EA., Fagot, J., Young, M.E. Same-different conceptualization by baboons (Papio papio): The role of entropy. Journal of Comparative Psychology 115, 2001, 42-52.

Katz, J.S., Wright, A.A. Same/different abstract-concept learning by pigeons. Journal of Experimental Psychology: Animal Behavior Processes 32, 2006, 80-86.

Wright, A.A.; Katz, J.S. Mechanisms of same/different concept learning in primates and avians. Behavioural Processes 72, 2006, 234-254.
Meeting 4: A switch to invertebrates: Cognition in spiders and bees? Lest we become too enamored of studies on animals we are sure are intelligent, let’s take a brief detour into the world of those whom we would assume are mostly reflexive critters. Having looked at categorization and same-different in vertebrates, can we now look at invertebrates as well? [Don’t worry about the gory details of the equipment….]
Jackson, R.R., Wilcox, R. S. Aggressive mimicry, prey-specific predatory behaviour and predator-recognition in the predator-prey interactions of Portia fimbriata and Euryattus sp., jumping spiders from Queensland. Behavioral Ecology and Sociobiology 26, 1990, 111-119.

Jackson, R.R., Wilcox, R.S. Spider flexibly chooses aggressive mimicry signals for different prey by trial and error. Behaviour 127, 1993, 21-36.

Wilcox, R.S., Jackson, R.R. Gentile, K. Spiderweb smokescreens: Spider trickster uses background noise to mask stalking movements. Animal Behaviour 51, 1996, 313-326.

Gould, J.L. Honey bee cognition. Cognition 37, 1990, 83-103.

Giurfa, M., Zhang, S., Jenett, A., Menzel, R., Srinivasan, M.V. The concepts of 'sameness' and 'difference' in an insect. Nature 410, 2001, 930-933.

Meeting 5: Number concepts: another cognitive task. Lenneberg once stated that animals will never succeed on tasks involving language or numerical concepts because they lacked abstraction and relational abilities. Well, animals likely aren’t going to show evidence of Fermat’s theorem or give the lecture for me, but they can understand numerical concepts.
Boysen, ST., Berntson, G.G. Numerical competence in a chimpanzee (Pan troglodytes). .
Journal of Comparative Psychology 103, 1989, 23-31.

Pepperberg, I.M. Numerical competence in an African Grey parrot. Journal of Comparative Psychology 108, 1994, 36 44.

Hauser, M.D., Carey, S. Spontaneous representations of small numbers of objects by rhesus macaques: Examinations of content and format. Cognitive Psychology 47, 2003, 367-401.

Tomonaga, M., Matsuzawa, T. Enumeration of briefly presented items by the chimpanzee (Pan troglodytes) and humans (Homo sapiens). Animal Learning & Behavior 30, 2002, 143-157.

Pepperberg, I.M., Gordon, J.D. Numerical comprehension by a Grey Parrot (Psittacus erithacus), including a zero-like concept. Journal of Comparative Psychology 119, 2005, 197-209.

Pepperberg, I.M. Grey Parrot (Psittacus erithacus) numerical abilities: addition and further experiments on a zero-like concept. Journal of Comparative Psychology 120, 2006, 1-11.

Pepperberg, I.M. Ordinality and inferential abilities of a Grey Parrot (Psittacus erithacus). Journal of Comparative Psychology 120, 2006, 205-216.

Cantlon, J.F.; Brannon, E. M. Shared system for ordering small and large numbers in monkeys and humans. Psychological Science 17, 2006, 401-406.


Meeting 6: Spatial concepts: caching behavior and a modular mind. Many animals cache food hoards, but birds have been studied most extensively. Birds who do cache seem to have specialized abilities that do not transfer to other behavior patterns. What can we learn about looking at a specialized system?
Shettleworth, S.J., Hampton, R.H. Adaptive specializations of spatial cognition in food storing birds? Approaches to testing a comparative hypothesis. In Animal Cognition in Nature, RP Balda, IM Pepperberg, AC Kamil, Eds., pp. 65-98, San Diego: Academic Press, 1998.

Shettleworth, SJ., Westwood, R.P. Divided attention, memory, and spatial discrimination in food-storing and nonstoring birds, black-capped chickadees (Poecile atricapilla) and dark-eyed Juncos (Junco hyemalis). Journal of Experimental Psychology: Animal Behavior Processes 28, 2002, 227-241.

Olson, DJ., Kamil, A.C., Balda, R.P., Nims, P.J. Performance of four-seed caching corvid species in operant tests of nonspatial and spatial memory. Journal of Comparative Psychology 109,1995, 173-181.

Bednekoff, P.A.., Balda, R.P., Kamil, A.C., Hile, Arla G. Long-term spatial memory in four seed-caching corvid species. Animal Behaviour 53,1997, 335-341.



Meeting 7: Caching and episodic memory: Scrub jays. Although the researchers have not proven that these birds have episodic memory…not just knowing what but also knowing when…their work comes closest to such proof. These birds also exhibit other types of awareness when caching.
de Kort, S.R., Dickinson, A., Clayton, N.S. Retrospective cognition by food-caching western scrub-jays. Learning and Motivation 36, 2005, 159-176.

Hampton, R.R., Hampstead, B.M., Murray, E.A Rhesus monkeys (Macaca mulatta) demonstrate robust memory for what and where, but not when, in an open-field test of memory. Learning and Motivation 36, 2005, 245-259.

Suddendorf, T., Busby, J. Like it or not? The mental time travel debate: Reply to Clayton et al. Trends in Cognitive Sciences 7, 2003, 437-438.

Emery, N.J., Dally, J.M., Clayton, N.S. Western scrub-jays (Aphelocoma californica) use cognitive strategies to protect their caches from thieving conspecifics.


Animal Cognition 7, 2004, 37-43.

Dally, J.M.; Emery, N.J.; Clayton, N.S. Food-caching western scrub-jays keep track of who was watching when. Science 312(5780), 2006, 1662-1665. [New!]



Meeting 8: Insightful behavior. Many studies, from early work of Kohler and chimpanzees, show some level of insightful behavior. Here we concentrate mostly on more recent work with birds.
Osthaus, B., Lea, S.E.G., Slater, A.M. Dogs (Canis lupus familiaris) fail to show understanding of means-end connections in a string-pulling task. Animal Cognition 8, 2005, 37-47.

Pepperberg, I.M."Insightful" string-pulling in Grey parrots (Psittacus erithacus) is affected by vocal competence. Animal Cognition 7, 2004, 263-266.

Heinrich, B. An experimental investigation of insight in Common Ravens (Corvus corax). Auk. 112, 1995, 994-1003.

Heinrich, B. Testing insight in ravens. In The Evolution of Cognition, C Heyes, L Huber, Eds., pp. 289-305. Cambridge, MA: MIT Press, 2000.

Heinrich, B., Bugnyar, T. Testing problem solving in ravens: string-pulling to reach food.
Ethology 111, 2005, 962-976.

Meeting 9: Imitative behavior. What exactly is imitation, and does it exist in animals? We will examine some differences among cultural transmission, emulation, and imitation and discuss animal ‘imitative’ behavior. We won’t be able to examine all important papers, but at least we will introduce the topic.
Whiten, A., Horner, V., Litchfield, C.A., Marshall-Pescini, S. How do apes ape? Learning & Behavior. 32, 2004, 36-52.

Caldwell, C.A., Whiten, A. Testing for social learning and imitation in common marmosets, Callithrix jacchus, using an artificial fruit. Animal Cognition 7, 2004, 77-85.

Gajdon, G.K., Fijn, N., Huber, L. Testing social learning in a wild mountain parrot, the kea (Nestor notabilis). Learning & Behavior 32, 2004, 62-71.

Voelkl, B., Huber, L. True imitation in marmosets. Animal Behaviour 60, 2000, 195-202.

Pepperberg, IM When parrots do not “parrot” (MS under review)

Meeting 10: Bird song: A cognitive process? Birds are thought to be pure mimics with respect to song…yes, they learn to sing, but then what? In the wild, birds actually learn not just what to sing, but the appropriate context in which to sing. Such behavior suggests cognitive processing
Kroodsma, D.E., Byers, B.E. Songbird song repertoires: an ethological approach to studying cognition. In R.P. Balda, I.M. Pepperberg, A.C. Kamil, Eds., Animal Cognition in Nature, pp. 305-336. London: Academic.1998.

Todt, D., Hultsch, H. Hierarchical learning, development and representation of song. In R. Balda, I.M. Pepperberg, A.C. Kamil, Eds., Animal Cognition in Nature, pp. 275-303. London: Academic, 1998.

Beecher, M.D, Campbell, S.E., Burt, J.M., Hill, C.E., and Nordby, J.C. Song-type matching between neighbouring song sparrows. Animal Behaviour 59, 2000, 21-27.

Burt, J.M., Bard, S.C., Campbell, S.E., and Beecher, M.D. 2002. Alternative forms of song matching in song sparrows. Animal Behaviour 63, 2002, 1143-1151.

Freeberg, T.M., King, A.P., West, M.E. Cultural transmission of vocal traditions in cowbirds (Molothrus ater) influences courtship patterns and mate preferences. Journal of Comparative Psychology 115, 2001, 201-211.
Meeting 11: Transitive inference: A strong test of cognition. In the wild, animals with a social dominance hierarchy likely understand transitive inference: e.g., if Sam beat up on Joe, and Joe beat up on me, then Sam is likely to smash me to smithereens. But demonstrating this behavior in the laboratory is not quite so simple.

Paz-y-Miño C.G. Bond, A.B.; Kamil, A.C.; Balda, R.P. Pinyon jays use transitive inference to predict social dominance. Nature 430, 2004, 778-781.

Bond, A.B., Kamil, A.C., Balda, R.P. Social complexity and transitive inference in corvids.
Animal Behaviour 65, 2003, 479-487.

Benard, J., Giurfa, M. A test of transitive inferences in free-flying honeybees: Unsuccessful performance due to memory constraints. Learning & Memory 11, 2004, 328-336.

Batsell, W. R. A classroom simulation of transitivity problems in animals. Teaching of Psychology 20, 1993, 228-230.

Boysen, S.T., Berntson, G.G., Shreyer, T.A., Quigley, K.S. Processing of ordinality and transitivity by chimpanzees (Pan troglodytes). Journal of Comparative Psychology 107, 1993, 208-215.

Gillan, D.J. Reasoning in the chimpanzee: II. Transitive inference. Journal of Experimental Psychology: Animal Behavior Processes 7, 1981, 150-164.

Meeting 12: Dolphin cognition. These are incredibly intelligent animals and we must examine their abilities…but, lacking hands and speech and being aquatic, these animals are more difficult to study than other species. Nevertheless, considerable information has arisen from studies on their capacities.
Sarko, D., Marino, L., Reiss, D.L. A bottlenose dolphin's (Tursiops truncatus) responses to its mirror image: Further analysis. International Journal of Comparative Psychology 15, 2002, 69-76.

Reiss, D., Marino, L. Mirror self-recognition in the bottlenose dolphin: a case of cognitive convergence. Proceedings of the National Academy of Sciences, USA 98, 2001, 5937-5942

Reiss, D., McCowan, B. Spontaneous vocal mimicry and production by bottlenose dolphins (Tursiops truncatus): Evidence for vocal learning. Journal of Comparative Psychology 107, 1993, 301-312.

Pack, A., Herman, L.M., Hoffmann-Kuhnt, M., Branstetter, B.K. The object behind the echo: Dolphins (Tursiops truncatus) perceive object shape globally through echolocation. Behavioural Processes 58, 2002, 1-26.

Herman, L.M., Kuczaj, S.A.; Holder, M.D. Responses to anomalous gestural sequences by a language-trained dolphin: Evidence for processing of semantic relations and syntactic information. Journal of Experimental Psychology: General 122, 1993, 184-194.

Meeting 13: A brief review of animal-language studies, Part I. We’ve touched on this work in other sections, but have not examined it in depth. An entire course would be necessary to understand the projects, their successes and their problems. But we’ll take a brief look at work with chimpanzees this week…[Yes, the reading list is really heavy, but there’s less for next week…..]
Jensvold, M.L.A.; Gardner, R.A. Interactive use of sign language by cross-fostered chimpanzees (Pan troglodytes). Journal of Comparative Psychology 114, 335-346.

Gardner, R.A., Gardner, B.T. A vocabulary test for chimpanzees (Pan troglodytes). Journal of Comparative Psychology 98, 1984, 381-404.

Premack, D. Possible general effects of language training on the chimpanzee. Human Development 27, 1984, 268-281.

Premack, D. Words: What are they, and do animals have them? Cognition 37, 1990, 197-212.

Brakke, K.E., Savage-Rumbaugh, E.S. The development of language skills in bonobo and chimpanzee: I. Comprehension. Language & Communication 15, 1995, 121-148.

Brakke, K.E., Savage-Rumbaugh, E.S. The development of language skills in pan: II. Production. Language & Communication 16,1996, 361-380.



Meeting 14: A brief review of animal-language studies, Part II. We’ll look again at the dolphin paper discussed earlier, then sea lions and parrots….
Pepperberg, I.M., Shive, H.A. Simultaneous development of vocal and physical object combinations by a Grey Parrot (Psittacus erithacus): Bottle caps, lids, and labels. Journal of Comparative Psychology 115, 2001, 376-384.

Pepperberg, I.M., Wilcox, S.E. Evidence for a form of mutual exclusivity during label acquisition by Grey parrots (Psittacus erithacus)? Journal of Comparative Psychology. 114, 2000, 219-231.

Gisiner, R., Schusterman, R. J. Sequence, syntax, and semantics: Responses of a language-trained sea lion (Zalophus californianus) to novel sign combinations. Journal of Comparative Psychology. 106, 1992, 78-91.

Schusterman, R.J., Kastak, D. California sea lion (Zalophus californianus) is capable of forming equivalence relations. Psychological Record 43, 1993, 823-839.



Meeting 15: Intelligence versus rationality and possibly consciousness? Does the first imply the other two? A review of several projects including more Grey parrot research. No readings for this lecture. [Remember, final projects are due next week!]
Meeting 16: Class projects are due. Class will be Q&A, discussing in detail any material from the class that the students wish to revisit.


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