Animal cognition - Knowledge

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targets; other birds and a number of other animals have been studied as well. In a typical experiment, a bird or other animal confronts a computer monitor on which a large number of pictures appear one by one, and the subject gets a reward for pecking or touching a picture of a category item and no reward for non-category items. Alternatively, a subject may be offered a choice between two or more pictures. Many experiments end with the presentation of items never seen before; successful sorting of these items shows that the animal has not simply learned many specific stimulus-response associations. A related method, sometimes used to study relational concepts, is matching-to-sample. In this task an animal sees one stimulus and then chooses between two or more alternatives, one of which is the same as the first; the animal is then rewarded for choosing the matching stimulus.

1186:. One individual called "Betty" spontaneously made a wire tool to solve a novel problem. She was being tested to see whether she would select a wire hook rather than a straight wire to pull a little bucket of meat out of a well. Betty tried poking the straight wire at the meat. After a series of failures with this direct approach, she withdrew the wire and began directing it at the bottom of the well, which was secured to its base with duct tape. The wire soon became stuck, whereupon Betty pulled it sideways, bending it and unsticking it. She then inserted the hook into the well and extracted the meat. In all but one of 10 subsequent trials with only straight wire provided, she also made and used a hook in the same manner, but not before trying the straight wire first. 1352:". Choosing "half empty" is supposed to indicate pessimism whereas choosing "half full" indicates optimism. To test this in animals, an individual is trained to anticipate that stimulus A, e.g. a 100 Hz tone, precedes a positive event, e.g. highly desired food is delivered when a lever is pressed by the animal. The same individual is trained to anticipate that stimulus B, e.g. a 900 Hz tone, precedes a negative event, e.g. bland food is delivered when the animal presses a lever. The animal is then tested by being given an intermediate stimulus C, e.g. a 500 Hz tone, and observing whether the animal presses the lever associated with the positive or negative reward. This has been suggested to indicate whether the animal is in a positive or negative mood. 1190:

associate sound and meaning, demolishing long-held theories that birds were only capable of mimicking human voices. Studies by other researchers have determined that African Grays can use deductive reasoning to correctly choose between pairs of boxes containing food and boxes that are empty. Until Pepperberg began this research in the 1970s, few scientists had studied intelligence in parrots, and few do today. Most inquiries have instead focused on monkeys, chimpanzees, gorillas, and dolphins, all of which are much more difficult to raise, feed, and handle. By the late 1980s, Alex had learned the names of more than 50 different objects, five shapes, and seven colors. He'd also learned what "same" and "different" mean—a step so crucial in human intellectual development

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animal responds consistently to A, a second stimulus ("B") accompanies A on additional training trials. Later tests with the B stimulus alone elicit little response, suggesting that learning about B has been blocked by prior learning about A. This result supports the hypothesis that stimuli are neglected if they fail to provide new information. Thus, in the experiment just cited, the animal failed to attend to B because B added no information to that supplied by A. If true, this interpretation is an important insight into attentional processing, but this conclusion remains uncertain because blocking and several related phenomena can be explained by models of conditioning that do not invoke attention.

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for pecking at pictures in set A but not for pecks at pictures in set B. After they had learned this task fairly well, the outcome was reversed: items in set B led to food and items in set A did not. Then the outcome was reversed again, and then again, and so on. Vaughan found that after 20 or more reversals, associating a reward with a few pictures in one set caused the birds to respond to the other pictures in that set without further reward as if they were thinking "if these pictures in set A bring food, the others in set A must also bring food." That is, the birds now categorized the pictures in each set as functionally equivalent. Several other procedures have yielded similar results.

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related to the stimulus in some other way. In Hunter's studies, for example, a light appeared briefly in one of three goal boxes and then later the animal chose among the boxes, finding food behind the one that had been lighted. Most research has been done with some variation of the "delayed matching-to-sample" task. For example, in the initial study with this task, a pigeon was presented with a flickering or steady light. Then, a few seconds later, two pecking keys were illuminated, one with a steady light and one with a flickering light. The bird got food if it pecked the key that matched the original stimulus.

1850:(1823–1915), setting the stage for all subsequent experiments of this kind, argued that insects "obey their compelling instinct, without realizing what they do". For instance, to understand that she can grab her paralyzed prey by a leg instead of an antenna is utterly beyond the powers of a sand wasp. "Her actions are like a series of echoes each awakening the next in a settled order, which allows none to sound until the previous one has sounded." Fabre's numerous experiments led him, in turn, to the view that scientists often try to "exalt animals" instead of objectively studying them. 1892:, the ladder of nature on which animals of different species occupy successively higher rungs, with humans typically at the top. However, there is some disagreement with the use of such a hierarchy, with some critics saying it may be necessary to understand specific cognitive capacities as adaptations to differing ecological niches. Some biologists argue that humans are not, in fact, the smartest animal, and that no animal can be characterized as the smartest, given that some animals have superior cognitive skills in certain areas. This contrasts with evolutionary psychologists such as 684:

general concepts. Better evidence is provided if, after training, an animal successfully makes a choice that matches a novel sample that it has never seen before. Monkeys and chimpanzees do learn to do this, as do pigeons if they are given a great deal of practice with many different stimuli. However, because the sample is presented first, successful matching might mean that the animal is simply choosing the most recently seen "familiar" item rather than the conceptually "same" item. A number of studies have attempted to distinguish these possibilities, with mixed results.

702:(2008) trained rats to discriminate between visual sequences. For one group ABA and BAB were rewarded, where A="bright light" and B="dim light". Other stimulus triplets were not rewarded. The rats learned the visual sequence, although both bright and dim lights were equally associated with reward. More importantly, in a second experiment with auditory stimuli, rats responded correctly to sequences of novel stimuli that were arranged in the same order as those previously learned. Similar sequence learning has been demonstrated in birds and other animals as well. 1856:'s (1852–1936) observations suggested to him that prima facie intelligent behavior in animals is often the result of either instincts or trial and error. For instance, most visitors watching Morgan's dog smoothly lifting a latch with the back of its head (and thereby opening a garden gate and escaping) were convinced that the dog's actions involved thinking. Morgan, however, carefully observed the dog's prior, random, purposeless actions and argued that they involved "continued trial and failure, until a happy effect is reached", rather than "methodical planning". 586:

pigeons gain a reward only by choosing the correct combination of the two stimuli (e.g. a high frequency tone together with a yellow light). The birds perform well at this task, presumably by dividing attention between the two stimuli. When only one of the stimuli varies and the other is presented at its rewarded value, discrimination improves on the variable stimulus but discrimination on the alternative stimulus worsens. These outcomes are consistent with the notion that attention is a limited resource that can be more or less focused among incoming stimuli.

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food in a box, the animal must return to the central platform. The maze may be used to test both reference and working memory. Suppose, for example, that over a number of sessions the same 4 arms of an 8-arm maze always lead to food. If in a later test session the animal goes to a box that has never been baited, this indicates a failure of reference memory. On the other hand, if the animal goes to a box that it has already emptied during the same test session, this indicates a failure of working memory. Various confounding factors, such as odor

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of the water. When placed in the tank, the animal swims around until it finds and climbs up on the platform. With practice, the animal finds the platform more and more quickly. Reference memory is assessed by removing the platform and observing the relative amount of time the animal spends swimming in the area where the platform had been located. Visual and other cues in and around the tank may be varied to assess the animal's reliance on landmarks and the geometric relations among them.

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phase. During the habituation phase the animal is placed in an empty test arena. This is followed by the adaptation phase, where the animal is placed in the arena with two identical objects. In the third phase, the test phase, the animal is placed in the arena with one of the familiar objects from the previous phase and with one novel object. Based on the rodents innate curiosity, the animals that remember the familiar object will spend more time on investigating the novel object.

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several alternatives. For example, several studies have shown that performance is better on, for example, a color discrimination (e.g. blue vs green) after the animal has learned another color discrimination (e.g. red vs orange) than it is after training on a different dimension such as an X shape versus an O shape. The reverse effect happens after training on forms. Thus, the earlier learning appears to affect which dimension, color or form, the animal will attend to.

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light, nearly every animal that has been studied has been shown to have a "biological clock" that yields cycles of activity even when the animal is in constant illumination or darkness. Circadian rhythms are so automatic and fundamental to living things – they occur even in plants – that they are usually discussed separately from cognitive processes, and the reader is referred to the main article (

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pecks at a picture showing a moth. Crucially, the probability with which a particular species of moth was detected was higher after repeated trials with that species (e.g. A, A, A,...) than it was after a mixture of trials (e.g. A, B, B, A, B, A, A...). These results suggest again that sequential encounters with an object can establish an attentional predisposition to see the object.

1735:, 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 599:

several types were available. Tinbergen suggested that this prey selection was caused by an attentional bias that improved detection of one type of insect while suppressing detection of others. This "attentional priming" is commonly said to result from a pretrial activation of a mental representation of the attended object, which Tinbergen called a "searching image".

1862:(1874–1949) placed hungry cats and dogs in enclosures "from which they could escape by some simple act, such as pulling at a loop of cord". Their behavior suggested to him that they did not "possess the power of rationality". Most books about animal behavior, Thorndike wrote, "do not give us a psychology, but rather a eulogy of animals". 442:, who argued that "mind" is simply a name for processes in the head that control complex behavior, and that it is 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". 1509:

dolphin when it chose the board with the fewer number of dots. In the experimental trials, two boards were set up, and the dolphin would emerge from the water and point to one board. The dolphins chose the arrays with fewer dots at a rate much larger than chance, indicating they can differentiate between quantities. A particular

1692:, which is widely regarded as having high cognitive abilities. These birds have been observed to hide their food when dominant ravens are visible and audible at the same time. Based on this observation, ravens were tested for their understanding of "seeing" as a mental state. In a first step, the birds protected their 1304:

bananas hung out of reach. Instead, they behaved in a manner that was "unwaveringly purposeful", spontaneously placing boxes so that they could climb to reach the fruit. Modern research has identified similar behavior in animals usually thought of as much less intelligent, if appropriate pre-training is given.

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Daniel J. Povinelli and Timothy Eddy of the University of Louisiana showed that young chimpanzees, when given a choice between two food providers, were just as likely to beg food from a person who could see the begging gesture as from a person who could not, thereby raising the possibility that young

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when dominants were visible but not when they could only be heard from an adjacent room. In the next step, they had access to a small peephole which allowed them to see into the adjacent room. With the peephole open, the ravens guarded their caches against discovery when they could hear dominants in

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In a study that used this approach, rats that were playfully tickled responded differently than rats that were simply handled. The rats that had been tickled were more optimistic than the handled rats. The authors suggested that they had demonstrated "...for the first time a link between the directly

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behavior. However, these behaviors are often quite inflexible and cannot be applied effectively in new situations. A great many species of birds build nests with a wide range of complexities, but although nest-building behaviour fulfills the criteria of some definitions of "tool-use", this is not the

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Whether an animal ranges over a territory measured in square kilometers or square meters, its survival typically depends on its ability to do such things as find a food source and then return to its nest. Sometimes such a task can be performed rather simply, for example by following a chemical trail.

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A commonly-used variation of the matching-to-sample task requires the animal to use the initial stimulus to control a later choice between different stimuli. For example, if the initial stimulus is a black circle, the animal learns to choose "red" after the delay; if it is a black square, the correct

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trained pigeons to respond to the presence or absence of human beings in photographs. The birds readily learned to peck photos that contained partial or full views of humans and to avoid pecking photos with no human, despite great differences in the form, size, and color of both the humans displayed

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Perceptual categorization is said to occur when a person or animal responds in a similar way to a range of stimuli that share common features. For example, a squirrel climbs a tree when it sees Rex, Shep, or Trixie, which suggests that it categorizes all three as something to avoid. This sorting of

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Another way to produce attentional priming in search is to provide an advance signal that is associated with the target. For example, if a person hears a song sparrow he or she may be predisposed to detect a song sparrow in a shrub, or among other birds. A number of experiments have reproduced this

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Attention is a limited resource and is not a none-or-all response: the more attention devoted to one aspect of the environment, the less is available for others. A number of experiments have studied this in animals. In one experiment, a tone and a light are presented simultaneously to pigeons. The

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Other experiments have shown that after animals have learned to respond to one aspect of the environment responsiveness to other aspects is suppressed. In "blocking", for example, an animal is conditioned to respond to one stimulus ("A") by pairing that stimulus with reward or punishment. After the

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have claimed special attention, but all sorts of animals large and small (birds, fish, ants, bees, and others) have been brought into the laboratory or observed in carefully controlled field studies. In the laboratory, animals push levers, pull strings, dig for food, swim in water mazes, or respond

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experiments are often cited as providing support for the animal cognition hypothesis, his book is replete with counterexamples. For instance, he placed chimpanzees in a situation where they could only get bananas by removing a box. The chimpanzee, Köhler observed, "has special difficulty in solving

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have demonstrated the ability to monitor their own mental states and use an "I don't know" response to avoid answering difficult questions. Unlike the mirror test, which reveals awareness of the condition of one's own body, this uncertainty monitoring is thought to reveal awareness of one's internal

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Experiments using the peak procedure and other methods have shown that animals can time short intervals quite exactly, can time more than one event at once, and can integrate time with spatial and other cues. Such tests have also been used for quantitative tests of theories of animal timing, such as

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Survival often depends on an animal's ability to time intervals. For example, rufous hummingbirds feed on the nectar of flowers, and they often return to the same flower, but only after the flower has had enough time to replenish its supply of nectar. In one experiment hummingbirds fed on artificial

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It has been hypothesized that animals such as apes and wolves are good at spatial cognition because this skill is necessary for survival. Some researchers argue that this ability may have diminished somewhat in dogs because humans have provided necessities such as food and shelter during some 15,000

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keeps track of its position in this way as it wanders for many meters searching for food. Though it travels in a randomly twisted path, it heads straight home when it finds food. However, if the ant is picked up and released some meters to the east, for example, it heads for a location displaced by

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When an animal is unable to see its goal, it may learn the appearance of nearby objects and use these landmarks as guides. Researchers working with birds and bees have demonstrated this by moving prominent objects in the vicinity of nest sites, causing returning foragers to hunt for their nest in a

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is used to test an animal's memory for spatial location and to discover how an animal is able to determine locations. Typically the maze is a circular tank filled with water that has been made milky so that it is opaque. Located somewhere in the maze is a small platform placed just below the surface

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Tinbergen's field observations on priming have been supported by a number of experiments. For example, Pietrewicz and Kamil (1977, 1979) presented blue jays with pictures of tree trunks upon which rested either a moth of species A, a moth of species B, or no moth at all. The birds were rewarded for

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As noted above, the function of attention is to select information that is of special use to the animal. Visual search typically calls for this sort of selection, and search tasks have been used extensively in both humans and animals to determine the characteristics of attentional selection and the

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Human and non-human animal cognition have much in common, and this is reflected in the research summarized below; most of the headings found here might also appear in an article on human cognition. Of course, research in the two also differs in important respects. Notably, much research with humans

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presses a lever for food. A light comes on, a lever-press brings a food pellet at a fixed later time, say 10 seconds, and then the light goes off. Timing is measured during occasional test trials on which no food is presented and the light stays on. On these test trials, the rat presses the lever

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The behavior of most animals is synchronized with the earth's daily light-dark cycle. Thus, many animals are active during the day, others are active at night, still others near dawn and dusk. Though one might think that these "circadian rhythms" are controlled simply by the presence or absence of

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is used to test memory for spatial location and to determine the mental processes by which location is determined. In a radial maze test, an animal is placed on a small platform from which paths lead in various directions to goal boxes; the animal finds food in one or more goal boxes. Having found

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Perceptually unrelated stimuli may come to be responded to as members of a class if they have a common use or lead to common consequences. An oft-cited study by Vaughan (1988) provides an example. Vaughan divided a large set of unrelated pictures into two arbitrary sets, A and B. Pigeons got food

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of "concept" was discussed for hundreds of years by philosophers before it became a focus of psychological study. Concepts enable humans and animals to organize the world into functional groups; the groups may be composed of perceptually similar objects or events, diverse things that have a common

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Animals process information from eyes, ears, and other sensory organs to perceive the environment. Perceptual processes have been studied in many species, with results that are often similar to those in humans. Equally interesting are those perceptual processes that differ from, or go beyond those

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Moty Nissani of Wayne State University trained Burmese logging elephants to lift a lid in order to retrieve food from a bucket. The lid was then placed on the ground alongside the bucket (where it no longer obstructed access to the food) while the treat was simultaneously placed inside the bucket.

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of their surroundings, meaning that they acquire and use information that enables them to compute how far and in what direction to go to get from one location to another. Such a map-like representation is thought to be used, for example, when an animal goes directly from one food source to another

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The novel object recognition (NOR) test is an animal behavior test that is primarily used to assess memory alterations in rodents. It is a simple behavioral test that is based on a rodents innate exploratory behavior. The test is divided into three phases: habituation, training/adaptation and test

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Delayed response tasks are often used to study short-term memory in animals. Introduced by Hunter (1913), a typical delayed response task presents an animal with a stimulus such as colored light, and after a short time interval the animal chooses among alternatives that match the stimulus, or are

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As in humans, research with animals distinguishes between "working" or "short-term" memory from "reference" or long-term memory. Tests of working memory evaluate memory for events that happened in the recent past, usually within the last few seconds or minutes. Tests of reference memory evaluate

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and in the non-human pictures. In follow-up studies, pigeons categorized other natural objects (e.g. trees) and after training they were able without reward to sort photos they had not seen before . Similar work has been done with natural auditory categories, for example, bird songs. Honeybees (

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Convergent evidence indicates that non-human animals have the neuroanatomical, neurochemical, and neurophysiological substrates of conscious states along with the capacity to exhibit intentional behaviors. Consequently, the weight of evidence indicates that humans are not unique in possessing the

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In one study, rhesus monkeys viewed visual displays containing, for example, 1, 2, 3, or 4 items of different sorts. They were trained to respond to them in several ways involving numerical ordering, for example touching "1" first, "2" second and so on. When tested with displays containing items

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Although tool use was long assumed to be a uniquely human trait, there is now much evidence that many animals use tools, including mammals, birds, fish, cephalopods and insects. Discussions of tool use often involve a debate about what constitutes a "tool", and they often consider the relation of

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This is one of the simplest tests for memory spanning a short time interval. The test compares an animal's response to a stimulus or event on one occasion to its response on a previous occasion. If the second response differs consistently from the first, the animal must have remembered something

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to pick a larger or smaller amount. During training, the bears were rewarded with food for a correct response. All bears performed better than what random error predicted on the trials with static, non-moving dots, indicating that they could differentiate between the two quantities. The bears

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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. Moreover, ants of some species can count up to 20 and add and subtract

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It is clear that animals of quite a range of species are capable of solving problems that appear to require abstract reasoning; Wolfgang Köhler's (1917) work with chimpanzees is a famous early example. He observed that chimpanzees did not use trial and error to solve problems such as retrieving

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Another bird that is highly studied for its intelligence is the African Gray Parrot. American animal behaviorist and psychologist Irene Pepperberg vindicated that African Grays possess cognitive abilities. Pepperberg used a bird named "Alex" in her trials and was able to prove that parrots could

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Some animals appear to have an advanced understanding of their spatial environment and will not take the most direct route if this confers an advantage to them. Some jumping spiders take an indirect route to prey rather than the most direct route, thereby indicating flexibility in behaviour and

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Many animals readily classify objects by perceived differences in form or color. For example, bees or pigeons quickly learn to choose any red object and reject any green object if red leads to reward and green does not. Seemingly much more difficult is an animal's ability to categorize natural

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Experimental research on visual search in animals was initially prompted by field observations published by Luc Tinbergen (1960). Tinbergen observed that birds are selective when foraging for insects. For example, he found that birds tended to catch the same type of insect repeatedly even though

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Animal ability to process and respond to stimuli is correlated with brain size. Small-brain animals tend to show simple behaviors that are less dependent on learning than those of large-brained animals. Vertebrates, particularly mammals, have larger brains and complex behavior that changes with

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Animals trained to discriminate between two stimuli, say black versus white, can be said to attend to the "brightness dimension", but this says little about whether this dimension is selected in preference to others. More enlightenment comes from experiments that allow the animal to choose from

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have shown the ability to choose an array with fewer dots compared to one with more dots. Experimenters set up two boards showing various numbers of dots in a poolside setup. The dolphins were initially trained to choose the board with the fewer number of dots. This was done by rewarding the

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When tested in a simple stimulus matching-to-sample task (described above) many animals readily learn specific item combinations, such as "touch red if the sample is red, touch green if the sample is green." But this does not demonstrate that they distinguish between "same" and "different" as

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Still other experiments have explored nature of stimulus factors that affect the speed and accuracy of visual search. For example, the time taken to find a single target increases as the number of items in the visual field increases. This rise in reaction time is steep if the distracters are

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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.

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Most work on animal concepts has been done with visual stimuli, which can easily be constructed and presented in great variety, but auditory and other stimuli have been used as well. Pigeons have been widely used, for they have excellent vision and are readily conditioned to respond to visual

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refers to mental processes that select relevant information, inhibit irrelevant information, and switch among these as the situation demands. Often the selective process is tuned before relevant information appears; such expectation makes for rapid selection of key stimuli when they become

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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 what appears to the naive human observer to be intelligent behavior may be strictly attributable to simple

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flowers that quickly emptied of nectar but were refilled at some fixed time (e.g. twenty minutes) later. The birds learned to come back to the flowers at about the right time, learning the refill rates of up to eight separate flowers and remembering how long ago they had visited each one.

476:, and many other matters. Studies often focus on the behavior of animals in their natural environments and discuss the putative function of the behavior for the propagation and survival of the species. These developments reflect an increased cross-fertilization from related fields such as 1525:. Given the choice between two groups of differing size, the angelfish will choose the larger of the two groups. This can be seen with a discrimination ratio of 2:1 or greater, such that, as long as one group has at least twice the fish as another group, it will join the larger one. 1277:

are capable of complex tasks, thus earning them the reputation of being among the smartest of invertebrates. For example, octopuses can open jars to get the contents inside and have remarkable ability to learn new skills from the moment they are born. Some cephalopods are known to use

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approaches to animal behavior were fallacious, and that people should only consider behaviour as, for example, rational, purposive or affectionate, if there is no other explanation in terms of the behaviours of more primitive life-forms to which we do not attribute those faculties.

1117:, particularly the great apes. The use of tools by primates is varied and includes hunting (mammals, invertebrates, fish), collecting honey, processing food (nuts, fruits, vegetables and seeds), collecting water, weapons and shelter. Research in 2007 shows that chimpanzees in the 437:

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

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in humans (40-50%). However, results from a recent meta-analysis suggest that the average correlation between performance scores on various cognitive tasks is only 0.18. Results from this study suggest that current evidence for general intelligence is weak in non-human animals.

1429:'s theory that animals must solve problems gradually, by trial and error. He said that Thorndike's animals could only use trial and error because the situation precluded other forms of problem solving. He provided chimps with a relatively unstructured situation, and he observed 1498:

choosing correctly in congruent (number of dots coincided with area of the dots) and incongruent (number of dots did not coincide with area of the dots) trials suggests that they were indeed choosing between quantities that appeared on the screen, not just a larger or smaller

1408:, many researchers agree that many animals can understand the meaning of individual words, and that some may understand simple sentences and syntactic variations, but there is little evidence that any animal can produce new strings of symbols that correspond to new sentences. 1216:) in 2009 by Giacomo Bernardi. The fish fans sand to unearth the bivalve, takes it into its mouth, swims several meters to a rock, which it then uses as an anvil by smashing the mollusc apart with sideward thrashes of the head. This behaviour has also been recorded in a 248:, have speculated about the presence or absence of the animal mind. These speculations led to many observations of animal behavior before modern science and testing were available. This ultimately resulted in the creation of multiple hypotheses about animal intelligence. 54: 769:, 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. 697:

in which the "rule" consists of the order in which a series of events occurs. Rule use is shown if the animal learns to discriminate different orders of events and transfers this discrimination to new events arranged in the same order. For example, Murphy

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available. A large body of research has explored the way attention and expectation affect the behavior of non-human animals, and much of this work suggests that attention operates in birds, mammals and reptiles in much the same way that it does in humans.

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function, relationships such as same versus different, or relations among relations such as analogies. Extensive discussions on these matters together with many references may be found in Shettleworth (2010) Wasserman and Zentall (2006) and in Zentall

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has harnessed her unique expertise in animal welfare and the ethical treatment of farm livestock to highlight underlying similarities between humans and other animals. From a methodological point of view, one of the main risks in this sort of work is

403:. 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 the idea that mental processes control behavior was the 1492:

demonstrated quantity differentiation abilities in a task with a computer screen. The bears were trained to touch a computer monitor with a paw or nose to choose a quantity of dots in one of two boxes on the screen. Each bear was trained with

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numbers within 5. This has been demonstrated using carefully crafted experiments based on measuring the time it takes for a scouting ant to pass the information to its team about the branch of an experimental maze on which food can be found.

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either studies or involves language, and much research with animals is related directly or indirectly to behaviors important to survival in natural settings. Following are summaries of some of the major areas of research in animal cognition.

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Many animals travel hundreds or thousands of miles in seasonal migrations or returns to breeding grounds. They may be guided by the Sun, the stars, the polarization of light, magnetic cues, olfactory cues, winds, or a combination of these.

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The use of rules has sometimes been considered an ability restricted to humans, but a number of experiments have shown evidence of simple rule learning in primates and also in other animals. Much of the evidence has come from studies of

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even though its previous experience has involved only travel between each source and home. Research in this area has also explored such topics as the use of geometric properties of the environment by rats and pigeons, and the ability of

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set the direction of much 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 differences between Thorndike's

911:, also known as "path integration", is the process of computing one's position by starting from a known location and keeping track of the distances and directions subsequently traveled. Classic experiments have shown that the 450:

The acceleration of research on animal cognition in the last 50 years or so has led to a rapid expansion in the variety of species studied and methods employed. The remarkable behavior of large-brained animals such as

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Delfour F, Marten K (April 2001). "Mirror image processing in three marine mammal species: killer whales (Orcinus orca), false killer whales (Pseudorca crassidens) and California sea lions (Zalophus californianus)".

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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

1747:, but not for monkeys. The mirror test has been criticized by 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 328:

In no case is an animal activity to be interpreted in terms of higher psychological processes if it can be fairly interpreted in terms of processes which stand lower in the scale of psychological evolution and

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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

1896:, who assess, based on the large list of related unique characteristics that humans do possess, that humans evolved to fill a unique "cognitive niche" and can fairly be characterized as the smartest animal. 646:

instances into groups is crucial to survival. Among other things, an animal must categorize if it is to apply learning about one object (e.g. Rex bit me) to new instances of that category (dogs may bite).

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such problems; he often draws into a situation the strangest and most distant tools, and adopts the most peculiar methods, rather than remove a simple obstacle which could be displaced with perfect ease".

410:

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,

1517:. After number and vocalization training, this was done by asking the parrot how many objects there were in a display. The parrot was able to identify the correct amount at a rate higher than chance. 3409:

Bhatt RS, Wasserman EA, Reynolds WF, Knauss KS (July 1988). "Conceptual behavior in pigeons: Categorization of both familiar and novel examples from four classes of natural and artificial stimuli".

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more and more until about 10 sec and then, when no food comes, gradually stops pressing. The time at which the rat presses most on these test trials is taken to be its estimate of the payoff time.

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placed humans and animals on a continuum, although Darwin's largely anecdotal approach to the cognition topic would not pass scientific muster later on. This method would be expanded by his protégé

1437:) were shown to exhibit similar insightful problem solving. A male was observed moving a box to a position where it could be stood upon to reach food that had been deliberately hung out of reach. 1786:

neurological substrates that generate consciousness. Non-human animals, including all mammals and birds, and many other creatures, including octopuses, also possess these neurological substrates.

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vertebrate. This has contributed to a common perception amongst researchers that mammals and birds share similar "advanced" cognitive characteristics as humans, while other vertebrates such as

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Typically, however, the animal must somehow acquire and use information about locations, directions, and distances. The following paragraphs outline some of the ways that animals do this.

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A variety of studies indicates that animals are able to use and communicate quantitative information, and that some can count in a rudimentary way. Some examples of this research follow.

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Wüstenberg D, Gerber B, Menzel R (August 1998). "Short communication: long- but not medium-term retention of olfactory memories in honeybees is impaired by actinomycin D and anisomycin".

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Several species of birds have been observed to use tools in the wild, including warblers, parrots, Egyptian vultures, brown-headed nuthatches, gulls and owls. Some species, such as the

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when hunting, considered the first evidence of systematic use of weapons in a species other than humans. Other mammals that spontaneously use tools in the wild or in captivity include

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of Skinner. This view seeks to explain behavior, including "private events" like mental images, solely by reference to the environmental contingencies impinging on the human or animal.

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In July, 2012 during the "Consciousness in Human and Nonhuman Animals" conference in Cambridge a group of scientists announced and signed a declaration with the following conclusions:

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are more "primitive", which has led to them being understudied. Despite this, increasing evidence indicates that fish possess not just capabilities that cannot be explained through

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Herrmann E, Call J, Hernàndez-Lloreda MV, Hare B, Tomasello M (September 2007). "Humans have evolved specialized skills of social cognition: the cultural intelligence hypothesis".

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given a choice between two displays with differing amounts of fruit flies, used as a food reward, reliably choose the display with more flies, as shown in a particular experiment.

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pick up stones and other small objects with their mandibles and drop them down the vertical entrances of rival colonies, allowing workers to forage for food without competition.

1286:. Cephalopod cognitive evolution is hypothesized to have been shaped primarily by predatory and foraging pressures, but a challenging mating context may also have played a role. 829:

choice is "green". Ingenious variations of this method have been used to explore many aspects of memory, including forgetting due to interference and memory for multiple items.

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conditioning alone, such as reversal learning, novel obstacle avoidance, and passing simultaneous two-choice tasks, but also even more complex capabilities such as navigational

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refers to a systematic pattern of deviation from norm or rationality in judgment, whereby inferences about other individuals or situations may be drawn in an illogical fashion.

7941:"Baboons can recognize written words, study finds; The monkeys don't assign meaning to them, but learn what letter combinations are common to real words, the study authors say" 1319:) from urbanized areas are better at innovative problem-solving tasks than bullfinches from rural environments, but that they did not differ in colour discrimination learning. 373:

associations. According to Thorndike, using Morgan's Canon, the inference of animal reason, insight, or consciousness is unnecessary and misleading. At about the same time,

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Anderson US, Stoinski TS, Bloomsmith MA, Marr MJ, Smith AD, Maple TL (August 2005). "Relative numerousness judgment and summation in young and old Western lowland gorillas".

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has been found to fit data from a sample of mammals. The formula is suggestive at best, and should only be applied to non-mammals with extreme caution. For some of the other

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they had never seen before, they continued to respond to them in order. The authors conclude that monkeys can represent the numerosities 1 to 9 at least on an ordinal scale.

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Frasnelli, Elisa; Robert, Théo; Chow, Pizza Ka Yee; Scales, Ben; Gibson, Sam; Manning, Nicola; Philippides, Andrew O.; Collett, Thomas S.; Hempel de Ibarra, Natalie (2020).

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given the choice between two food trays demonstrated the ability to choose the tray with more food items at a rate higher than chance after training. In a similar task,

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Carazo P, Font E, Forteza-Behrendt E, Desfilis E (May 2009). "Quantity discrimination in Tenebrio molitor: evidence of numerosity discrimination in an invertebrate?".

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The details of interval timing have been studied in a number of species. One of the most common methods is the "peak procedure". In a typical experiment, a rat in an

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Jaakkola K, Fellner W, Erb L, Rodriguez M, Guarino E (August 2005). "Understanding of the concept of numerically "less" by bottlenose dolphins (Tursiops truncatus)".

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insightful changes of behavior, as, for example, when a chimp suddenly moved a box into position so that it could retrieve a banana. More recently, Asian elephants (

9915: 8215:

Akinrinade, Ibukun; Kareklas, Kyriacos; Teles, Magda C.; Reis, Thais K.; Gliksberg, Michael; Petri, Giovanni; Levkowitz, Gil; Oliveira, Rui F. (March 23, 2023).

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Walraven V, van Elsacker L, Verheyen R (1995). "Reactions of a group of pygmy chimpanzees (Pan paniscus) to their mirror images: evidence of self-recognition".

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Early inferences were not necessarily precise or accurate. Nonetheless, interest in animal mental abilities, and comparisons to humans, increased with early

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Original was Intelligenzprüfungen an Anthropoiden, Berlin 1917. 2nd German edition was titled Intelligenzprüfungen an Menschenaffen, Berlin: Springer 1921.

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Gómez-Laplaza LM, Gerlai R (January 2011). "Can angelfish (Pterophyllum scalare) count? Discrimination between different shoal sizes follows Weber's law".

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Animals often learn what their nest or other goal looks like, and if it is within sight they may simply move toward it; it is said to serve as a "beacon".

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published research on artificial language comprehension in the bottlenose dolphin. Though this sort of research has been controversial, especially among

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Rushton JP (2004). "Placing intelligence into an evolutionary framework or how g fits into the r–K matrix of life-history traits including longevity".

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by Weir (1962) and in investigations of early speech in children by Greenfield and others (1976). Some such research has been done with a macaw (see

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and its refinement over the years. Still, Romanes is most famous for two major flaws in his work: his focus on anecdotal observations and entrenched

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Bräuer J, Kaminski J, Riedel J, Call J, Tomasello M (February 2006). "Making inferences about the location of hidden food: social dog, causal ape".

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Several experiments cannot be readily reconciled with the belief that some animal species are intelligent, insightful, or possess a theory of mind.

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to put coins into a box. The raccoon drifted to its instinctive behavior of rubbing the coins with its paws, as it would do when foraging for food.

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Some researchers propose that animal calls and other vocal behaviors provide evidence of consciousness. This idea arose from research on children's

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similar to the target, less steep if they are dissimilar, and may not occur if the distracters are very different from the target in form or color.

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Dooling RJ, Okanoya K (1995). "Psychophysical methods for assessing perceptual categories.". In Klump GM, Dooling RJ, Fay RR, Stebbins WC (eds.).

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There is some evidence for cognitive bias in a number of species, including rats, dogs, rhesus macaques, sheep, chicks, starlings and honeybees.

261:, in which a crow drops pebbles into a vessel of water until he is able to drink. This was a relatively accurate reflection of the capability of 1899:

Whether fairly or not, the performance of animals is often compared to that of humans on cognitive tasks. Our closest biological relatives, the

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All elephants continued to toss the lid before retrieving the reward, thus suggesting that elephants do not grasp simple causal relationships.

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where an animal's sense organs transmitted information to an organ capable of making decisions, and then to a motor organ. Despite Aristotle's

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Blough DS (2006). "Reaction-time explorations of visual attention, perception, and decision in pigeons.". In Wasserman EA, Zentall TR (eds.).

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traveling with shells collected for protection. Despite evolving independently from humans for over 600 million years, octopuses demonstrate

12429: 762:, whose ecological niches require them to remember the locations of thousands of caches, often following radical changes in the environment. 7512: 5120:

Möglich MH, Alpert GD (1979). "Stone dropping by Conomyrma bicolor (Hymenoptera: Formicidae): A new technique of interference competition".

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is usually credited with introducing the concept of insight into experimental psychology. Working with chimpanzees, Köhler came to dispute

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studies have shown that a single factor of intelligence is responsible for 47% of the individual variance in cognitive ability measures in

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Uller C, Jaeger R, Guidry G, Martin C (June 2003). "Salamanders ( Plethodon cinereus) go for more: rudiments of number in an amphibian".

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Wright AA, Santiago HC, Sands SF, Kendrick DF, Cook RG (July 1985). "Memory processing of serial lists by pigeons, monkeys, and people".

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in non-human animals cannot be measured with verbal scales, it has been measured using a variety of methods that involve such things as

1025:("SET"), Killeen's Behavioral Theory of Timing, and Machado's Learning to Time model. No one theory has yet gained unanimous agreement. 12434: 10937: 8667: 11350: 4417:

Savolainen P, Zhang YP, Luo J, Lundeberg J, Leitner T (November 2002). "Genetic evidence for an East Asian origin of domestic dogs".

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Campbell CB, Hodos W (September 1991). "The Scala naturae revisited: evolutionary scales and anagenesis in comparative psychology".

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Patterson FG, Cohn RH (1994). "Self-recognition and self-awareness in lowland gorillas". In Parker ST, Mitchell R, Boccia M (eds.).

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Tinbergen L (1960). "The natural control of insects in pine woods: I. Factors influencing the intensity of predation by songbirds".

1681:, to oneself and others and to understand that others have desires, intentions, and perspectives that are different from one's own. 472:

experiments. Careful field studies explore memory for food caches, navigation by stars, communication, tool use, identification of

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Brown MJ, Gordon DM (2000). "How resources and encounters affect the distribution of foraging activity in a seed-harvesting ants".

1400:

and his analysis of others results, Herbert Terrace criticized the idea that chimps can produce new sentences. Shortly thereafter

12895: 9314: 9216: 726:) have been applied to the study of animal memory, and some of the phenomena characteristic of human short term memory (e.g. the 2402: 9391: 7902:"Baboons can learn to recognize words; Monkeys' ability suggests that reading taps into general systems of pattern recognition" 5219:

Tebbich S, Seed AM, Emery NJ, Clayton NS (April 2007). "Non-tool-using rooks, Corvus frugilegus, solve the trap-tube problem".

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have been trained to use tools in controlled experiments. One species examined extensively under laboratory conditions is the

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Kundey SM, Strandell B, Mathis H, Rowan JD (2010). "Learning of monotonic and nonmonotonic sequences in domesticated horses (

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Fiset S, Plourde V (May 2013). "Object permanence in domestic dogs (Canis lupus familiaris) and gray wolves (Canis lupus)".

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Locurto C, Scanlon C (1998). "Individual differences and a spatial learning factor in two strains of mice (Mus musculus)".

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Blough DS, Blough PM (1990). "Reaction-time assessments of visual processes in pigeons.". In Berkley M, Stebbins W (eds.).

943:

when an animal or human searches their environment for specific objects to focus on among other objects in the environment.

7961: 2378:"Crows understand water displacement at the level of a small child: Show causal understanding of a 5- to 7-year-old child" 1767:, although this interpretation has been questioned. These species might also be aware of the strength of their memories. 11762: 9967: 8637: 8556: 8537: 8044:

Rodríguez, Fernando; Quintero, Blanca; Amores, Lucas; Madrid, David; Salas-Peña, Carmen; Salas, Cosme (August 11, 2021).

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Kamil AC, Bond AB (2006). "Selective attention, priming, and foraging behavior.". In Wasserman EA, Zentall TR (eds.).

11955: 11431: 11391: 11314: 11271: 11125: 11050: 10524: 9279: 9269: 9147: 8481: 8402: 6215: 2412: 1798:

can influence the interpretation of cognitive research. Instinctive drift is the tendency of an animal to revert to

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about the first, unless some other factor such as motivation, sensory sensitivity, or the test stimulus has changed.

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The mind and behavior of non-human animals has captivated the human imagination for centuries. Many writers, such as

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Locurto C, Durkin E. "Problem-solving and individual differences in mice (Mus musculus) using water reinforcement".

12524: 11599: 11250: 11080: 10907: 9977: 9972: 9402: 9079: 7849: 8363: 7979:

Vinepinsky, Ehud; Cohen, Lear; Perchik, Shay; Ben-Shahar, Ohad; Donchin, Opher; Segev, Ronen (September 8, 2020).

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Call JE, Burghardt GM, Pepperberg IM, Snowdon CT, Zentall TE, eds. (2017). "Chapter 23 : Timing in Animals".

4270: 3438:"Acoustic and perceptual categories of vocal elements in the warble song of budgerigars (Melopsittacus undulatus)" 2144:"Horse sense: social status of horses (Equus caballus) affects their likelihood of copying other horses' behavior" 12770: 11567: 11320: 11205: 10414: 10000: 9895: 8830: 7682: 2018: 1488:

Other experiments have been conducted that show animals' abilities to differentiate between non-food quantities.

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have also been shown to exhibit a number of higher-level skills such as tool use, but the amount of research on

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Mather JA, Kuba MJ (May 2013). "The cephalopod specialties: complex nervous systems, learning and cognition".

6352:"How general is cognitive ability in non-human animals? A meta-analytical and multi-level reanalysis approach" 788:

workers have more carrying capacity and thus more reason to memorise that information, and so they do. Slugs,

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objects that vary a great deal in color and form even while belonging to the same group. In a classic study,

7490:. Illustrated by Bob Walters (2nd ed.). Bloomington, Ind.: Indiana University Press. pp. 191–208. 4759: 4703:"Testing the Scalar Expectancy Theory (SET) and the Learning to Time model (LeT) in a double bisection task" 2923:

Kamin LJ (1969). "Predictability, surprise, attention, and conditioning". In Campbell BA, Church RM (eds.).

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provides an example of evidence for theory of mind in a non-human species. Ravens are members of the family

12880: 12472: 12168: 11282: 11240: 11060: 10454: 9385: 9330: 7901: 1965:, a 2019 review of evidence indicates they can experience numerous emotions, such as pleasure and anxiety. 1566: 1100: 936: 861: 3597: 2250: 1604:

construct that summarizes the correlations observed between an individual's scores on various measures of

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in fish, and extraordinary visual acuity, motion sensitivity and ability to see ultraviolet light in some

12517: 12250: 11738: 11527: 11155: 11105: 9789: 9264: 9209: 7981:"Representation of edges, head direction, and swimming kinematics in the brain of freely-navigating fish" 5005:"Bipedal locomotion in Octopus vulgaris: A complementary observation and some preliminary considerations" 4032:"Behavioral analysis of internal memory states using cooling-induced retrograde amnesia in Limax flavus" 738:; some of this work has sought to clarify the physiological basis of spatial memory and the role of the 12451: 12200: 11980: 11852: 11647: 11583: 11519: 11170: 11130: 11100: 10228: 10223: 9625: 7427: 5763:

Reznikova, Zhanna; Ryabko, Boris (2011). "Numerical competence in animals, with an insight from ants".

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Herrnstein RJ (1979). "Acquisition, Generalization, and Discrimination Reversal of a Natural Concept".

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workers vary in their effort investment towards memorising flower locations, with smaller workers less

12343: 7165: 5171: 287:(mistaken belief that cognition occurred in the heart), this approached some modern understandings of 12875: 12195: 11964: 11802: 11639: 11075: 10369: 10288: 10148: 9728: 9058: 9033: 8932: 8705: 8653: 4976: 1927: 1657: 1617: 1531:

have been shown to be capable of numeracy, and some species can distinguish among numbers up to six.

867: 5777: 1513:, after training, has shown the ability to differentiate between the numbers zero through six using 12808: 12465: 12148: 11933: 11894: 11880: 11746: 11722: 11511: 11135: 10539: 10484: 9941: 9660: 9645: 9630: 9587: 9552: 9289: 9023: 8904: 8899: 7516: 2028: 1819: 1758:

in some animals provides some evidence for cognitive self-awareness. The great apes, dolphins, and

1594: 1586: 1393: 1212:(scallops, urchins and clams) shells. This behavior was first filmed in an orange-dotted tuskfish ( 257: 11866: 8272:"Given the Cold Shoulder: A Review of the Scientific Literature for Evidence of Reptile Sentience" 12890: 12792: 12123: 12015: 11786: 11690: 11503: 11479: 11356: 11235: 11190: 11085: 10972: 10781: 9457: 9375: 9096: 8800: 7520: 7270: 5956:"Size matters: impact of item size and quantity on array choice by chimpanzees (Pan troglodytes)" 5313:"The town bird and the country bird: problem solving and immunocompetence vary with urbanization" 2470: 2377: 2013: 1912: 1474: 1446: 1249: 485: 208: 97: 8422: 7962:"Baboons show their word skills; Reading may stem from a visual aptitude shared by all primates" 6350:

Poirier, Marc-Antoine; Kozlovsky, Dovid Y.; Morand-Ferron, Julie; Careau, Vincent (2020-12-09).

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Reznikova ZI (1999). "Ethological mechanisms of population dynamic in species ant communities".

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D'Amato MR, Colombo M (April 1988). "Representation of serial order in monkeys (Cebus apella)".

298:, the study of ant behavior, as well as the classification of humans as primates beginning with 12885: 12775: 12556: 12282: 12245: 12235: 12180: 12128: 12073: 12043: 11988: 11535: 11110: 11070: 11024: 10942: 10897: 10766: 10419: 10404: 10058: 9635: 9254: 9202: 9178: 9125: 8755: 8750: 8740: 6718:

Bischof-Köhler D (1991). "The development of empathy in infants". In Lamb ME, Keller H (eds.).

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Terrace HS, Petitto LA, Sanders RJ, Bever TG (November 1979). "Can an ape create a sentence?".

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Haselton MG, Nettle D, Andrews PW (2005). "The evolution of cognitive bias". In Buss DM (ed.).

2003: 1939: 1621: 1558: 1540: 1522: 1521:, when put in an unfamiliar environment will group together with conspecifics, an action named 1236:) in an aquarium setting. These species are at opposite ends of the phylogenetic tree in this 727: 396: 347: 321: 276: 85: 8365:

Animal Spatial Cognition: Comparative, Neural, and Computational Approaches. [On-line]

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Animal Spatial Cognition: Comparative, Neural, and Computational Approaches. [On-line]

4170:"Morris water maze: procedures for assessing spatial and related forms of learning and memory" 801:

memory for regularities such as "pressing a lever brings food" or "children give me peanuts".

11794: 11615: 11575: 11487: 11447: 11368: 11362: 11175: 10459: 10359: 10324: 9910: 9809: 9718: 9557: 9447: 9412: 9053: 9003: 8878: 8805: 8715: 2053: 1866: 1625: 1405: 1356:

measured positive affective state and decision making under uncertainty in an animal model".

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Bräuer, Juliane; Hanus, Daniel; Pika, Simone; Gray, Russell; Uomini, Natalie (2 July 2020).

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Nissani M (2005). "Do Asian elephants apply causal reasoning to tool use tasks? 31: 91–96".

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was a significant contribution to subsequent cognitive research in both humans and animals.

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Gatti RC (2015). "Self-consciousness: beyond the looking-glass and what dogs found there".

7089: 6986: 6803: 6674: 6494: 5585: 5484: 5414: 5357: 5076: 5016: 4790: 4623: 4426: 3932: 3694: 3612: 3332: 3142: 3087: 3005: 2311: 1993: 1988: 1943: 1888: 1799: 1706: 1232: 432: 392: 168: 120: 12398: 10686: 4219:"The novel object recognition memory: neurobiology, test procedure, and its modifications" 2251:"Social learning across species: horses (Equus caballus) learn from humans by observation" 8: 12729: 12724: 12648: 12479: 12260: 11912: 11859: 11276: 10887: 10499: 10494: 10374: 10349: 10068: 9925: 9814: 9708: 9703: 9173: 8868: 8863: 8783: 8745: 8571: 6038:"Bears "Count" Too: Quantity Estimation and Comparison in Black Bears (Ursus Americanus)" 4827:

Shettleworth SJ (December 2012). "Do animals have insight, and what is insight anyway?".

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Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences

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Much of what is happening in the world at any moment is irrelevant to current behavior.

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Sherwin CM (2001). "Can invertebrates suffer? Or, how robust is argument-by-analogy?".

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was the earliest to attest that said story reflects the behavior of real-life corvids.

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are beginning to clarify the physiological substrate of some inferred mental process.

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The common chimpanzee can use tools. This individual is using a stick to get food.

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