Animal cognition

Washoe, a female chimpanzee who was the first non-human to learn to communicate using American Sign Language, as part of a research experiment on animal language acquisition.

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.^{[citation needed]}

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 invertebrates such as cephalopods, spiders, and insects.^{[citation needed]}

Historical Background

Animal cognition from anecdote to laboratory

The behavior of non-human animals has captivated human imagination from antiquity, and over the centuries many writers have speculated about the animal mind, or its absence, as Descartes would have it. Speculation about animal intelligence gradually yielded to scientific study after Darwin placed humans and animals on a continuum, although Darwin’s largely anecdotal approach to the topic would not pass scientific muster later on. Unsatisfied with the anecdotal method of Darwin and his protégé Romanes, E. L. Thorndike brought animal behavior into the laboratory for objective scrutiny. Thorndike’s careful observations of the escape of cats, dogs, and chicks from puzzle boxes led him to conclude that intelligent behavior may be compounded of simple associations and that inference to animal reason, insight, or consciousness is unnecessary and misleading. At about the same time, I. P. Pavlov began his seminal studies of conditioned reflexes in dogs. Pavlov quickly abandoned attempts to infer canine mental processes; such attempts, he said, led only to disagreement and confusion. He was, however, willing to propose unseen physiological processes that might explain his observations.

The behavioristic half-century

The work of Thorndike and Pavlov, and a little later of the outspoken behaviorist John B. Watson set the direction of most research on animal behavior for more than half a century. During this time there was considerable progress in understanding simple associations; notably, around 1930 the difference between Thorndike’s instrumental (or operant) conditioning and Pavlov’s classical (or Pavlovian) conditioning was clarified, first by Miller and Kanorski and then by B. F. Skinner.,. Many experiments on conditioning followed; they generated some complex theories, but they made little or no reference to intervening mental processes. Probably the most explicit dismissal of “mental events” was the radical behaviorism of B. F. Skinner, which sought to explain behavior solely by reference to the environmental contingencies impinging on the human or animal.

Despite the predominantly behaviorist orientation of research before 1960, the rejection of mental processes in animals was not universal during those years. Influential exceptions included, for example, Wolfgang Köhler and his insightful chimpanzees and Edward Tolman whose proposed cognitive map was a significant contribution to subsequent cognitive research in both humans and animals.

The cognitive revolution

Beginning around 1960, a “cognitive revolution” in research on humans gradually spurred a similar transformation of research with animals. Inference to processes not directly observable became acceptable and then commonplace. An important proponent of this shift in thinking was Donald O. Hebb, who argued that “mind” is simply a name for processes in the head that control complex behavior, and that it both necessary and possible to infer those processes from behavior Animals came to be seen as “goal seeking agents that acquire, store, retrieve, and internally process information at many levels of cognitive complexity.” . However, it is interesting to note that may cognitive experiments with animals made, and still make, ingenious use of conditioning methods pioneered by Thorndike and Pavlov.

The scientific status of “consciousness” in animals continues to be hotly debated. Serious consideration of conscious thought in animals has been advocated by some (e.g.Donald Griffin), but the larger research community has been notably cool to such suggestions

Methods

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.^{[citation needed]}

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.^{[citation needed]}

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.^{[citation needed]}

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.^{[citation needed]} However, progress in the different areas has been variable. Among the fields of interest are:

Attention

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).^{[citation needed]}

Categorization

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.^{[citation needed]}

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

Memory

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.

Memory has been widely investigated in foraging honeybees, Apis mellifera, which use both transient short-term working memory that is non-feeder specific and a feeder specific long-term reference memory. Memory induced in a free-flying honeybee by a single learning trial lasts for days and, by three learning trials, for a lifetime. Slugs, Limax flavus, have a short-term memory of approximately 1 min and long-term memory of 1 month.

Spatial cognition

The ability to properly navigate and search through the environment is a critical task for many animals. Much of this movement seems to be directed, in the sense that the animals in question seem to be purposely moving towards a particular spot for a reason. Purposeful navigation implies some sort of cognitive map of the external environment. Research in this area (Brown & Cook, 2006) 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.^{[citation needed]}

Tool and weapon use

Further information: Tool use by animals

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.

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.

Some cephalopods are known to use coconut shells for protection or camouflage (see undersea video).^{[citation needed]}

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.”

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; 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.

Language

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.^{[citation needed]}

Consciousness

Mirror test with a baboon

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, 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. 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.^{[citation needed]}

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.^{[citation needed]}

Mathematics

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. Young chimpanzees have outperformed human college students in tasks requiring remembering numbers.

Ants are able to use quantitative values and transmit this information. For instance, ants of several species are able to estimate quite precisely numbers of encounters with members of other colonies on their feeding territories. Numeracy has been described in the yellow mealworm beetle, Tenebrio molitor, and the honeybee.

Cognitive faculty by species

Some animals such as dogs, horses, great apes, and (more recently) dolphins and parrots are typically thought by laymen 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.

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.

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, but the amount of research on cephalopod intelligence is still limited.^{[citation needed]}

See also

• Anthropomorphism
• Dog intelligence
• Pain in invertebrates#Cognitive abilities

Further reading

• Brown, M.F., & Cook, R.G. (Eds.). (2006). Animal Spatial Cognition: Comparative, Neural, and Computational Approaches. [On-line]. Available: www.pigeon.psy.tufts.edu/asc/
• 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

• The limits of intelligence Douglas Fox, Scientific American, 14 June 2011.
• Animal Cognition entry by Kristin Andrews in the Stanford Encyclopedia of Philosophy
• Animal Consciousness entry by Colin Allen in the Stanford Encyclopedia of Philosophy
• Animal Cognition Network
• Center for Avian Cognition University of Nebraska (Alan Kamil, Alan Bond)