Numerical cognition is a subdiscipline of cognitive science that studies the cognitive, developmental and neural bases of numbers and mathematics. As with many cognitive science endeavors, this is a highly interdisciplinary topic, and includes researchers in cognitive psychology, developmental psychology, neuroscience and cognitive linguistics. This discipline, although it may interact with questions in the philosophy of mathematics, is primarily concerned with empirical questions.

Topics included in the domain of numerical cognition include: How do non-human animals process numerosity? How do infants acquire an understanding of numbers (and how much is inborn)? How do humans associate linguistic symbols with numerical quantities? How do these capacities underlie our ability to perform complex calculations? What are the neural bases of these abilities, both in humans and in non-humans? What metaphorical capacities and processes allow us to extend our numerical understanding into complex domains such as the concept of infinity, the infinitesimal or the concept of the limit in calculus? Heuristics in numerical cognition

Comparative studies A variety of research has demonstrated that non-human animals, including rats, lions and various species of primates have an approximate sense of number (referred to as "numerosity"). For example, when a rat is trained to press a bar 8 or 16 times to receive a food reward, the number of bar presses will approximate a Gaussian or Normal distribution with peak around 8 or 16 bar presses. When rats are more hungry, their bar-pressing behavior is more rapid, so by showing that the peak number of bar presses is the same for either well-fed or hungry rats, it is possible to disentangle time and number of bar presses. In addition, in a few species the parallel individuation system has been shown, for example in the case of guppies which successfully discriminated between 1 and 4 other individuals.

Similarly, researchers have set up hidden speakers in the African savannah to test natural (untrained) behavior in lions. These speakers can play a number of lion calls, from 1 to 5. If a single lioness hears, for example, three calls from unknown lions, she will leave, while if she is with four of her sisters, they will go and explore. This suggests that not only can lions tell when they are "outnumbered" but that they can do this on the basis of signals from different sensory modalities, suggesting that numerosity is a multisensory concept.

Developmental studies Developmental psychology studies have shown that human infants, like non-human animals, have an approximate sense of number. For example, in one study, infants were repeatedly presented with arrays of (in one block) 16 dots. Careful controls were in place to eliminate information from "non-numerical" parameters such as total surface area, luminance, circumference, and so on. After the infants had been presented with many displays containing 16 items, they habituated, or stopped looking as long at the display. Infants were then presented with a display containing 8 items, and they looked longer at the novel display.

Because of the numerous controls that were in place to rule out non-numerical factors, the experimenters infer that six-month-old infants are sensitive to differences between 8 and 16. Subsequent experiments, using similar methodologies showed that 6-month-old infants can discriminate numbers differing by a 2:1 ratio (8 vs. 16 or 16 vs. 32) but not by a 3:2 ratio (8 vs. 12 or 16 vs. 24). However, 10-month-old infants succeed both at the 2:1 and the 3:2 ratio, suggesting an increased sensitivity to numerosity differences with age.

In another series of studies, Karen Wynn showed that infants as young as five months are able to do very simple additions (e.g., 1 + 1 = 2) and subtractions (3 - 1 = 2). To demonstrate this, Wynn used a "violation of expectation" paradigm, in which infants were shown (for example) one Mickey Mouse doll going behind a screen, followed by another. If, when the screen was lowered, infants were presented with only one Mickey (the "impossible event") they looked longer than if they were shown two Mickeys (the "possible" event). Further studies by Karen Wynn and Koleen McCrink found that although infants' ability to compute exact outcomes only holds over small numbers, infants can compute approximate outcomes of larger addition and subtraction events (e.g., "5+5" and "10-5" events).

There is debate about how much these infant systems actually contain in terms of number concepts, harkening to the classic nature versus nurture debate. suggested that a child innately has the concept of natural number, and only has to map this onto the words used in her language. disagreed, saying that these systems can only encode large numbers in an approximate way, where language-based natural numbers can be exact. Without language, only numbers 1 to 4 are believed to have an exact representation, through the parallel individuation system. One promising approach is to see if cultures that lack number words can deal with natural numbers. The results so far are mixed (e.g., ); , ).

Neuroimaging and neurophysiological studies Human neuroimaging studies have demonstrated that regions of the parietal lobe, including the intraparietal sulcus (IPS) and the inferior parietal lobule (IPL) are activated when subjects are asked to perform calculation tasks. Based on both human neuroimaging and neuropsychology, Stanislas Dehaene and colleagues have suggested that these two parietal structures play complementary roles. The IPS is thought to house the circuitry that is fundamentally involved in numerical estimation, number comparison, and on-line calculation, or quantity processing (often tested with subtraction) while the IPL is thought to be involved in rote memorization, such as multiplication. Thus, a patient with a lesion to the IPL may be able to subtract, but not multiply, and vice versa for a patient with a lesion to the IPS. In addition to these parietal regions, regions of the frontal lobe are also active in calculation tasks. These activations overlap with regions involved in language processing such as Broca's area and regions involved in working memory and attention. Additionally, the inferotemporal cortex is implicated in processing the numerical shapes and symbols, necessary for calculations with Arabic digits. More current research has highlighted the networks involved with multiplication and subtraction tasks. Multiplication is often learned through rote memorization and verbal repetitions, and neuroimaging studies have shown that multiplication uses a left lateralized network of the inferior frontal cortex and the superior-middle temporal gyri in addition to the IPL and IPS. Subtraction is taught more with quantity manipulation and strategy use, more reliant upon the right IPS and the posterior parietal lobule.

Single-unit neurophysiology in monkeys has also found neurons in the frontal cortex and in the intraparietal sulcus that respond to numbers. Andreas Nieder trained monkeys to perform a "delayed match-to-sample" task. For example, a monkey might be presented with a field of four dots, and is required to keep that in memory after the display is taken away. Then, after a delay period of several seconds, a second display is presented. If the number on the second display match that from the first, the monkey has to release a lever. If it is different, the monkey has to hold the lever. Neural activity recorded during the delay period showed that neurons in the intraparietal sulcus and the frontal cortex had a "preferred numerosity", exactly as predicted by behavioral studies. That is, a certain number might fire strongly for four, but less strongly for three or five, and even less for two or six. Thus, we say that these neurons were "tuned" for specific quantities. Note that these neuronal responses followed Weber's law, as has been demonstrated for other sensory dimensions, and consistent with the ratio dependence observed for non-human animals' and infants' numerical behavior.

While primates have remarkably similar brains to humans, there are differences in function, ability, and sophistication. They make for good preliminary test subjects, but do not show small differences that are the result of different evolutionary tracks and environment. However, in the realm of number, they share many similarities. As identified in monkeys, neurons selectively tuned to number were identified in the bilateral intraparietal sulci and prefrontal cortex in humans. Piazza and colleagues investigated this using fMRI, presenting participants with sets of dots where they either had to make same-different judgments or larger-smaller judgments. The sets of dots consisted of base numbers 16 and 32 dots with ratios in 1.25, 1.5, and 2. Deviant numbers were included in some trials in larger or smaller amounts than the base numbers. Participants displayed similar activation patterns as Nieder found in the monkeys. The intraparietal sulcus and the prefrontal cortex, also implicated in number, communicate in approximating number and it was found in both species that the parietal neurons of the IPS had short firing latencies, whereas the frontal neurons had longer firing latencies. This supports the notion that number is first processed in the IPS and, if needed, is then transferred to the associated frontal neurons in the prefrontal cortex for further numerations and applications. Humans displayed Gaussian curves in the tuning curves of approximate magnitude. This aligned with monkeys, displaying a similarly structured mechanism in both species with classic Gaussian curves relative to the increasingly deviant numbers with 16 and 32 as well as habituation. The results followed Weber's Law, with accuracy decreasing as the ratio between numbers became smaller. This supports the findings made by Nieder in macaque monkeys and shows definitive evidence for an approximate number logarithmic scale in humans.