Animal cognition

Animal cognition

Animal cognition is the title given to the study of the mental capacities of non-human animals. It has developed out of comparative psychology, but has also been strongly influenced by the approach of ethology, behavioral ecology, and evolutionary psychology. The alternative name cognitive ethology is therefore sometimes used; much of what used to be considered under the title of animal intelligence is now thought of under this heading.

In practice, animal cognition mostly concerns mammals, especially primates, cetaceans, and elephants, as well as dogs, cats, and rodents. However, research also extends to non-mammalian vertebrates such as birds including parrots, corvids, and pigeons, as well as lizards, snakes, and fish, even to non-vertebrates such as cephalopods, spiders, and insects.


Historical background

For most of the twentieth century, the dominant approach to animal psychology was to use experiments on intelligence in animals to uncover simple learning processes (such as classical conditioning and operant conditioning) that might then account for the apparently more complex intellectual abilities of humans. This approach is well summarized in the mid-century book by Hilgard (1958), but its reductionist philosophy was combined with a strongly behaviorist methodology, in which overt behavior was taken as the only valid data for the study of psychology, and in its more extreme forms (the radical behaviorism of B. F. Skinner and his experimental analysis of behavior) behavior was taken as the only topic of interest. In effect, the mental processes that humans experience in themselves were viewed as epiphenomena (see, for example, Skinner, 1969).

The success of cognitive psychology in addressing human mental processes, which began in the late 1950s and was proclaimed by Neisser (1967), led to a re-evaluation of the research paradigm, and researchers began to address animal mental processes from the opposite direction, by taking what is known about human mental processes and looking for evidence of comparable processes in other species. In a sense this was a return to the approach of Darwin's protégé George Romanes (e.g. 1886), arguably the first comparative psychologist of the modern era. However, whereas Romanes relied heavily on anecdote and an anthropomorphic projection of human capacities onto other species, modern researchers in animal cognition are in most cases firmly behaviorist in methodology, even though they differ sharply from the behaviorist philosophy.

There are some exceptions to the rule of behaviorist methodology, such as John Lilly and, some would argue, Donald Griffin (e.g. 1992), who have been prepared to take a strong position that other animals do have minds and that humans should approach the study of their cognition accordingly. However, their claims have not found wide acceptance in the scientific community, though they have attracted an enthusiastic following among lay people.

The development of animal cognition was also strongly influenced by:

  • increased use of and interest in primates (and also cetaceans) rather than the rats and pigeons that had become the classic species of the comparative psychology laboratory, and by developments within primatology;
  • advancing knowledge of animals' behavior in their natural environments through studies in ethology, sociobiology and behavioral ecology; such studies often showed that animals needed certain cognitive abilities in order to adapt to their ecological niche (as for example in studies of caching birds such as Clark's Nutcracker,[1]) or appeared to use cognitive abilities under natural conditions (for example in Jane Goodall's studies of chimpanzees, see Goodall (1991);
  • one or two high profile projects, in particular Allen and Beatrice Gardner's Washoe project in which a chimpanzee learned at least some elements of American Sign Language.
  • advancing understanding of brain function through work in physiological psychology and cognitive neuropsychology

This account of the history of the study of animal cognition is inevitably oversimplified. From Romanes on, there have always been comparative psychologists who have been more or less cognitively inclined: obvious examples are Wolfgang Köhler, famous for his studies of insight in chimpanzees, and Edward C. Tolman, who introduced into psychology, as an explanation of the behavior of rats in mazes, two ideas that have been immensely influential in human cognitive psychology - the cognitive map and the idea of decision-making in risky choice according to expected value.


Research in animal cognition continues to use some of the established research techniques of comparative psychology and the experimental analysis of behavior, such as mazes and Skinner boxes, though it employs them in new varieties (such as the 8-arm maze and Morris water maze that have been used in many studies of spatial memory) and in new ways. However, it complements those with observation of animals in their natural environments, or quasi-natural environments and also with field experiments.

It has also been characterized by a number of very long term projects, such as the Washoe project and other ape-language experiments (e.g. project Nim), Irene Pepperberg's extended series of studies with the African Gray Parrot Alex, Louis Herman's work with bottlenosed dolphins, and studies of long-term memory in pigeons in which birds were shown to remember pictures for periods of several years. Some cognitive research also requires the management of animal behavior, and the use of operant conditioning to facilitate animal training. In general, the conclusion of concept formation in an animal requires a generalization test where the animal responds appropriately to a novel stimulus to which associative learning cannot explain the response behavior.

Some researchers have made effective use of a Piagetian methodology, taking tasks which human children are known to master at different stages of development, and investigating which of them can be performed by particular species. Others have been inspired by concerns for animal welfare and the management of domestic species: for example Temple Grandin has harnessed her unique expertise in animal welfare and the ethical treatment of farm livestock to highlight underlying similarities between humans and other animals.

Research questions

The common Chimpanzee can use tools. This chimpanzee is using a stick in order to get food.

Given the broad program of animal cognition, the areas of study in animal cognition follow more or less from those in human cognitive psychology. However, progress in the different areas has been variable. Among the fields of interest are:


Research has focused on animals' ability to distribute attention between different aspects of a stimulus, and on visual search. As in humans, it appears that sharing attention between stimulus features reduces the capacity to detect any one of them, though there are some ecologically relevant visual search tasks at which particular species show remarkable abilities (for example, pigeons have an extraordinary capacity to pick out grain from substrate).


Following pioneering research by Richard Herrnstein, there has been a mass of research on birds' ability to discriminate between categories of stimuli, including the kinds of ill-defined category that are used in everyday human speech. Birds have been found to learn this kind of task easily, and to transfer correct responses readily to new instances of the categories.

It has also been found that rhesus monkeys understand same-different relationships,[2] easily form categories based on prototype theory,[3] and may even have some capacity for rule-based learning.


The categories that have been developed to analyze human memory (short term memory, long term memory, working memory) have been applied to the study of animal memory, and some of the phenomena characteristic of human short term memory (e.g. the serial position effect) have been detected in animals, particularly monkeys[citation needed]. However most progress has been made in the analysis of spatial memory, partly in relation to studies of the physiological basis of spatial memory and the role of the hippocampus, and partly in relation to scatter-hoarder animals such as Clark's Nutcracker, certain jays, tits and certain squirrels, whose ecological niches require them to remember the locations of thousands of caches, often[citation needed] following radical[citation needed] changes in the environment.

Spatial cognition

The ability to properly navigate and search through the environment is a critical task for many animals. Research in this area (Brown & Cook, 2006[4]) has focused on such diffuse topics as landmark and beacon use by ants and bees, the encoding and use of geometric properties of the environment by pigeons, and the ability of rats to represent a spatial pattern in either radial arm mazes or pole box mazes. Sometimes included under the envelope of spatial cognition is work in humans and other animals in visual search tasks, which aim to experimentally address questions about searching through one's environment for a particular object.

Tool and weapon use

Some species, such as the Woodpecker Finch of the Galapagos Islands, use particular tools as an essential part of their foraging behavior. However, these behaviors are often quite inflexible and cannot be applied effectively in new situations. Several species have now been shown to be capable of more flexible tool use. A well known example is Jane Goodall's observation of chimpanzees "fishing" for termites in their natural environment, and captive great apes are often observed to use tools effectively; several species of corvids have also been trained to use tools in controlled experiments, or use bread crumbs for bait-fishing [1].

Research in 2007 shows that chimpanzees in the Fongoli savannah sharpen sticks to use as spears when hunting, considered the first evidence of systematic use of weapons in a species other than humans.[5]

Some cephalopods are known to use coconut shells for protection or camouflage (see undersea video).

Reasoning and problem solving

Closely related to tool use is the study of reasoning and problem solving. It has been observed that the manner in which chimpanzees solve problems, such as that of retrieving bananas positioned out of reach, is not through trial-and-error. Instead, they were observed to proceed in a manner that was “unwaveringly purposeful.”[6]

It is clear that animals of quite a range of species are capable of solving a range of problems that are argued to involve abstract reasoning;[7] modern research has tended to show that the performances of Wolfgang Köhler's chimpanzees, who could achieve spontaneous solutions to problems without training, were by no means unique to that species, and that apparently similar behavior can be found in animals usually thought of as much less intelligent, if appropriate training is given.[citation needed] Causal reasoning has also been observed in rooks and New Caledonian crows.[8][9]


The modeling of human language in animals is known as animal language research. In addition to the ape-language experiments mentioned above, there have also been more or less successful attempts to teach language or language-like behavior to some non-primate species, including parrots and Great Spotted Woodpeckers. Louis Herman published research on artificial language comprehension in the bottlenosed dolphin using cognitive research methods at the height of the skepticism produced by Herbert Terrace's criticism of chimpanzee language experiments through his own results with the animal Nim Chimpsky. In particular, the focus on the comprehension mode only allowed cognitive methods of utilizing blinded observers to grade the animals' gross physical behavior, rather than trying to interpret putative language production. Herman's results (Herman, Richards, & Wolz, 1984) were published in the journal Cognition, regarding work on the dolphins Akeakamai and Phoenix. All such research has been controversial among cognitive linguists.


The sense in which animals can be said to have consciousness or a self-concept has been hotly debated; it is often referred to as the debate over animal minds. The best known research technique in this area is the mirror test devised by Gordon G. Gallup, in which an animal's skin is marked in some way while it is asleep or sedated, and it is then allowed to see its reflection in a mirror; if the animal spontaneously directs grooming behavior towards the mark, that is taken as an indication that it is aware of itself. Self-awareness, by this criterion, has been reported for chimpanzees and also for other great apes, the European Magpie,[10] some cetaceans and a solitary elephant, but not for monkeys. The mirror test has attracted controversy among some researchers because it is entirely focused on vision, the primary sense in humans, while other species rely more heavily on other senses such as the olfactory sense in dogs.[citation needed]

It has been suggested that metacognition in some animals provides some evidence for cognitive self-awareness.[11] The great apes, dolphins, and rhesus monkeys have demonstrated the ability to monitor their own mental states and use an "I don't know" response to avoid answering difficult questions. These species might also be aware of the strength of their memories. Unlike the mirror test, which relies primarily on body images and bodily self-awareness, uncertainty monitoring paradigms are focused on the kinds of mental states that might be linked to mental self-awareness.

A different approach to determine whether a non-human animal is conscious derives from passive speech research with a macaw (see Arielle). Some researchers propose that by passively listening to an animal's voluntary speech, it is possible to learn about the thoughts of another creature and to determine that the speaker is conscious. This type of research was originally used to investigate a child's crib speech by Weir (1962) and in investigations of early speech in children by Greenfield and others (1976). With speech-capable birds, the methods of passive-speech research open a new avenue for investigation.


Some animals are capable of distinguishing between different amounts and rudimentary counting. Elephants have been known to perform simple arithmetic, and rhesus monkeys can count.[12][13] Ants are able to use quantitative values and transmit this information.[14][15] For instance, ants of several species are able to estimate quite precisely numbers of encounters with members of other colonies on their feeding territories.[16][17] Young chimpanzees have outperformed human college students in tasks requiring remembering numbers.[18] Pigeons have been shown to outperform humans on the Monty Hall problem, a probability puzzle.[19]

Cognitive faculty by species

Some animals such as dogs, horses, great apes, and (more recently) dolphins and parrots are typically thought by laymen[clarification needed] as intelligent in ways that some other species of animal are not.[citation needed] For example, crows are attributed with human-like intelligence in the folklore of many cultures. A number of recent survey studies have demonstrated the consistency of these rankings between people in a given culture and indeed to a considerable extent across cultures.[20]

A common image is the scala naturae, the ladder of nature on which animals of different species occupy successively higher rungs, with humans typically at the top.[21]

A more fruitful approach has been to recognize that different animals may have different kinds of cognitive processes, which are better understood in terms of the ways in which they are cognitively adapted to their different ecological niches, than by positing any kind of hierarchy. (See Shettleworth (1998), Reznikova (2007).)

One question that can be asked coherently is how far different species are intelligent in the same ways as humans are, i.e., are their cognitive processes similar to ours. Not surprisingly, our closest biological relatives, the great apes, tend to do best on such an assessment. Among the birds, corvids and parrots have typically been found to perform well. Despite ambitious claims, evidence of unusually high human-like intelligence among cetaceans is patchy, partly because the cost and difficulty of carrying out research with marine mammals mean that experiments frequently suffer from small sample sizes and inadequate controls and replication.[citation needed] Octopodes have also been shown to exhibit a number of higher-level skills such as tool use,[22] but the amount of research on cephalopod intelligence is still limited.

See also

Nicolas P. Rougier's rendering of the human brain.png Thinking portal


  1. ^ *Kamil, A. C.; Balda, R. P. (1990). "Differential memory for different cache sites by Clark's nutcrackers (Nucifraga columbiana)". Journal of Experimental Psychology: Animal Behavior Processes 16 (2): 162–168. doi:10.1037/0097-7403.16.2.162. 
  2. ^ Smith, J. D.; Redford, J. S., Coutinho, M. V. C., Couchman, J. J. (2008). "The comparative psychology of same-different judgments by humans (Homo sapiens) and monkeys (Macaca mulatta)". Journal of Experimental Psychology: Animal Behavior Processes 34 (2): 361–374. doi:10.1037/0097-7403.34.3.361. PMID 18665719. 
  3. ^ Couchman, J. J.; Coutinho, M. V. C., Smith, J. D. (2010). "Rules and Resemblance: Their Changing Balance in the Category Learning of Humans (Homo sapiens) and Monkeys (Macaca mulatta)". Journal of Experimental Psychology: Animal Behavior Processes 36 (2): 172–183. doi:10.1037/a0016748. PMC 2890302. PMID 20384398. 
  4. ^ Animal Spatial Cognition:Comparative, Neural & Computational Approaches
  5. ^ Chimps Use "Spears" to Hunt Mammals, Study Says John Roach for National Geographic News (February 22, 2007) (accessed on June 12, 2010)
  6. ^ Wolfgang Köhler The Mentality of Apes (1917)
  7. ^ For Chimpanzees, see for example David Premack (1983) The Mind of an Ape
  8. ^ "Non-tool-using rooks, Corvus frugilegus, solve the trap-tube problem". Anim Cogn 10 (2): 225–31. April 2007. doi:10.1007/s10071-006-0061-4. PMID 17171360. 
  9. ^ doi:10.1098/rspb.2008.1107
  10. ^ Prior, Schwarz, and Güntürkün, Helmut, Ariane, and Onur; Schwarz, A; Güntürkün, O; De Waal, Frans (2008). De Waal, Frans. ed. "Mirror-Induced Behavior in the Magpie (Pica pica): Evidence of Self-Recognition". PLoS Biology (Public Library of Science) 6 (8): e202. doi:10.1371/journal.pbio.0060202. PMC 2517622. PMID 18715117. Retrieved 2008-08-21. 
  11. ^ Couchman, Justin J.; Coutinho, M. V. C., Beran, M. J., & Smith, J. D. (2010). "Beyond Stimulus Cues and Reinforcement Signals: A New Approach to Animal Metacognition". Journal of Comparative Psychology 124 (4): , 356 –368. doi:10.1037/a0020129. 
  12. ^ Elephants show flair for arithmetic
  13. ^ Representation of the Numerosities 1-9 by Rhesus Macaques (Macaca mulatto
  14. ^ Zhanna Reznikova, Boris Ryabko, "A Study of Ants' Numerical Competence". Electronic Transactions on Artificial Intelligence, Issue: Vol. 5(2001): Section B: pp. 111-126
  15. ^ Reznikova, Zh. I. (2007). Animal Intelligence: From Individual to Social Cognition. Cambridge University Press
  16. ^ Reznikova, Zh. I. (1999). Ethological mechanisms of population dynamic in species ant communities. Russian Journal of Ecology, 30, 3, 187–197
  17. ^ Brown, M. J. F., Gordon, D. M. (2000). How resources and encounters affect the distribution of foraging activity in a seed-harvesting ants. Behav.Ecol.Sociobiol. , 47, 195-203.
  18. ^ Rowan Hooper (2007-12-03). "Chimps outperform humans at memory task". New Scientist. Retrieved 2008-03-24. 
  19. ^ doi:10.1037/a0017703
  20. ^ *Nakajima, S.; Arimitsu, K.; Lattal, K. M. (2002). "Estimation of animal intelligence by university students in Japan and the United States". Anthrozoös 15 (3): 194–205. doi:10.2752/089279302786992504. 
  21. ^ Campbell, C.B.G., & Hodos, W. (1991). The Scala Naturae revisited: Evolutionary scales and anagenesis in comparative psychology. J. Comp. Psychol. 105:211-221
  22. ^ Finn, J. K., Tregenza, T., & Norman M.D. (2009). Defensive tool use in a coconut-carrying octopus. Current Biology. DOI:10.1016/j.cub.2009.10.052

Further reading

  • Brown, M.F., & Cook, R.G. (Eds.). (2006). Animal Spatial Cognition: Comparative, Neural, and Computational Approaches. [On-line]. Available:
  • Goodall, J. (1991). Through a window. London: Penguin.
  • Griffin, D. R. (1992). Animal minds. Chicago: University of Chicago Press.
  • Hilgard, E. R. (1958). Theories of learning, 2nd edn. London: Methuen.
  • Neisser, U. (1967). Cognitive psychology. New York, Appleton-Century-Crofts.
  • Romanes, G. J. (1886). Animal intelligence, 4th edn. London: Kegan Paul, Trench.
  • Shettleworth, S. J. (1998). Cognition, evolution and behavior. New York: Oxford University Press.
  • Skinner, B. F. (1969). Contingencies of reinforcement: a theoretical analysis. New York: Appleton-Century-Crofts.
  • Narby, Jeremy. (2005) Intelligence In Nature. New York: Penguin.
  • Lurz, Robert W. (2009) Mindreading Animals: The Debate over What Animals Know about Other Minds. The MIT Press.

External links

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