Number sense in human infants

Four experiments used a preferential looking method to investigate 6-month-old infants' capacity to represent numerosity in visual-spatial displays. Building on previous findings that such infants discriminate between arrays of eight versus 16 discs, but not eight versus 12 discs (Xu & Spelke, 2000), Experiments 1 and 2 investigated whether infants' numerosity discrimination depends on the ratio of the two set sizes with even larger numerosities. Infants successfully discriminated between arrays of 16 versus 32 discs, but not 16 versus 24 discs, providing evidence that their discrimination shows the set-size ratio signature of numerosity discrimination in human adults, children and many non-human animals. Experiments 3 and 4 addressed a controversy concerning infants' ability to discriminate large numerosities (observed under conditions that control for total filled area, array size and density, item size and correlated properties such as brightness: Brannon, 2002; Xu, 2003b; Xu & Spelke, 2000) versus small numerosities (not observed under conditions that control for total contour length: Clearfield & Mix, 1999). To investigate the sources of these differing findings, Experiment 3 tested infants' large-number discrimination with controls for contour length, and Experiment 4 tested small-number discrimination with controls for total filled area. Infants successfully discriminated the large-number displays but showed no evidence of discriminating the small-number displays. These findings provide evidence that infants have robust abilities to represent large numerosities. In contrast, infants may fail to represent small numerosities in visual-spatial arrays with continuous quantity controls, consistent with the thesis that separate systems serve to represent large versus small numerosities.     

Small and large number processing in infants and toddlers with Williams syndrome

Previous studies have suggested that typically developing 6‐month‐old infants are able to discriminate between small and large numerosities. However, discrimination between small numerosities in young infants is only possible when variables continuous with number (e.g. area or circumference) are confounded. In contrast, large number discrimination is successful even when variables continuous with number are systematically controlled for. These findings suggest the existence of different systems underlying small and large number processing in infancy. How do these develop in atypical syndromes? Williams syndrome (WS) is a rare neurocognitive developmental disorder in which numerical cognition has been found to be impaired in older children and adults. Do impairments of number processing have their origins in infancy? Here this question is investigated by testing the small and large number discrimination abilities of infants and toddlers with WS. While infants with WS were able to discriminate between 2 and 3 elements when total area was confounded with numerosity, the same infants did not discriminate between 8 and 16 elements, when number was not confounded with continuous variables. These findings suggest that a system for tracking the features of small numbers of object (object‐file representation) may be functional in WS, while large number discrimination is impaired from an early age onwards. Finally, we argue that individual differences in large number processing in infancy are more likely than small number processing to be predictive of later development of numerical cognition.     

Do fish count? Spontaneous discrimination of quantity in female mosquitofish

The spontaneous tendency to join the largest social group was used to investigate quantity discrimination in fish. Fish discriminated between shoals that differed by one element when the paired numbers were 1vs2, 2vs3 and 3vs4, but not when 4vs5 or larger. Using large numerosities (>4), the ability to discriminate between two numbers improved as the numerical distance between them increased and a significant discrimination was found only with ratios of 1:2 or smaller (4vs8, 8vs16 and 4vs10). Experiments to control for non-numerical variables evidenced the role played by the total area of stimuli with both large and small numerosities; the total quantity of movement of the fish within a shoal appeared also important but only when large numerosities were involved. Even though the pattern of discrimination exhibited by female mosquitofish is not fully consistent with any of the existing models of quantity representation, our results seem to suggest two distinct mechanisms in fish, one used to compare small numbers of objects and one used when larger numerosities are involved.     

Quantity discrimination in Tenebrio molitor: evidence of numerosity discrimination in an invertebrate?

Numerosity discrimination, the ability to distinguish between sets with more and less items, is recognised as the foundation for higher numerical abilities. Understanding numerosity discrimination from a comparative perspective is hence pivotal in tracing the evolution of numerical representation systems. However, numerosity discrimination has been well studied only in vertebrates, where two innate systems of number representation have been described: an ‘analog magnitude system’ used to discriminate among numerosities by representing them as cardinal magnitudes and a ‘parallel individualisation system’ that allows precise discrimination among small arrays of items (≤4) by representing objects individually. We investigated the existence of quantity discrimination in an insect species (Tenebrio molitor) by using a spontaneous two-choice procedure in which males were exposed to substrates bearing odours from different numbers of females (≤4) in increasing numerosity ratios (1:4, 1:3 and 1:2). We show that males can discriminate sources of odours reflecting 1 versus 4 and 1 versus 3 females, but not 2 versus 4 or 1 versus 2, indicating that T. molitor males exhibit a marked preference for sources reflecting more female donors only when numerosity ratios are below 1:2. We discuss the functional significance of this finding and whether our pattern of results could be best explained by summation of a non-numerical continuous variable or by the existence of a numerosity discrimination mechanism with an operational signature ratio of 1:2.     

Do infants have a sense of numerosity? A p‐curve analysis of infant numerosity discrimination studies

Research demonstrating that infants discriminate between small (e.g., 1 vs. 3 dots) and large numerosities (e.g., 8 vs. 16 dots) is central to theories concerning the origins of human numerical abilities. To date, there has been no quantitative meta‐analysis of the infant numerical competency data. Here, we quantitatively synthesize the evidential value of the available literature on infant numerosity discrimination using a meta‐analytic tool called p‐curve. In p‐curve the distribution of available p‐values is analyzed to determine whether the published literature examining particular hypotheses contains evidential value. p‐curves demonstrated evidential value for the hypotheses that infants can discriminate between both small and large unimodal and cross‐modal numerosities. However, the analyses also revealed that the published data on infants’ ability to discriminate between large numerosities is less robust and statistically powered than the data on their ability to discriminate small numerosities. We argue there is a need for adequately powered replication studies to enable stronger inferences in order to use infant data to ground theories concerning the ontogenesis of numerical cognition.     

Auditory-visual intermodal matching of small numerosities in 6-month-old infants

Recent studies have reported that preverbal infants are able to discriminate between numerosities of sets presented within a particular modality. There is still debate, however, over whether they are able to perform intermodal numerosity matching, i.e. to relate numerosities of sets presented with different sensory modalities. The present study investigated auditory-visual intermodal matching of small numerosities in infancy by using a violation-of-expectation paradigm. After being familiarized with events of a few objects impacting a surface successively, 6-month-old infants were alternatively presented with two and three tones while the movement of each object remained hidden behind an opaque screen. The screen was then removed to reveal either two or three objects. Results showed that the infants looked significantly longer at the numerically nonequivalent events (the three-tone/two-object and the two-tone/three-object events) than at the numerically equivalent events (the two-tone/two-object and the three-tone/three-object events) irrespective of the rate or duration of auditory tones presented. These findings suggest that infants are capable of performing intermodal matching of small numerosities and that they might possess abstract representations of numerosity beyond sensory modalities.     

Dissociation between small and large numerosities in newborn infants

In the first year of life, infants possess two cognitive systems encoding numerical information: one for processing the numerosity of sets of 4 or more items, and the second for tracking up to 3 objects in parallel. While a previous study showed the former system to be already present a few hours after birth, it is unknown whether the latter system is functional at this age. Here, we adapt the auditory‐visual matching paradigm that previously revealed sensitivity to large numerosities to test sensitivity to numerosities spanning the range from 2 to 12. Across studies, newborns discriminated pairs of large numerosities in a 3:1 ratio, even when the smaller numerosity was 3 (3 vs. 9). In contrast, newborn infants failed to discriminate pairs including the numerosity 2, even at the same ratio (2 vs. 6). These findings mirror the dissociation that has been reported with older infants, albeit with a discontinuity situated between numerosities 2 and 3. Two alternative explanations are compatible with our results: either newborn infants have a separate system for processing small sets, and the capacity of this system is limited to 2 objects; or newborn infants possess only one system to represent numerosities, and this system either is not functional or is extremely imprecise when it is applied to small numerosities.     

Imprinted numbers: newborn chicks’ sensitivity to number vs. continuous extent of objects they have been reared with

Newborn chicks were tested for their sensitivity to number vs. continuous physical extent of artificial objects they had been reared with soon after hatching. Because of the imprinting process, such objects were treated by chicks as social companions. We found that when the objects were similar, chicks faced with choices between 1 vs. 2 or 2 vs. 3 objects chose the set of objects of larger numerosity, irrespective of the number of objects they had been reared with. Moreover, when volume, surface or contour length were controlled for using sets of 1 vs. 4, 1 vs. 6 or 1 vs. 3 objects, chicks resorted to choosing the larger object, rather than the familiar numerosity. When, however, chicks were reared with objects differing in their aspect (colour, size, and shape) and then tested with completely novel objects (of different colour and shape but controlled for continuous extent), they chose to associate with the same number of objects they had been reared with. These results suggest that identification of objects as different and separate individuals is crucial for the computation of number rather than continuous extent in numerical representation of small numerosities and provide a striking parallel with results obtained in human infants. Early availability of small numerosity discrimination by chicks strongly suggests that these abilities are in place at birth.     

Spontaneous discrimination of small quantities: shoaling preferences in angelfish (<Emphasis Type="Italic">Pterophyllum scalare</Emphasis>)

The ability to quantify, i.e. to estimate quantity, may provide evolutionary advantages in some contexts and has been demonstrated in a variety of animal species. In a prior study, we showed that angelfish (Pterophyllum scalare) were able to discriminate between groups (shoals) in which a large number of conspecifics swam preferring to join the larger of the two. Our results implied that angelfish can compare relative shoal sizes likely on the basis of some quantitative attributes of the shoal. Here, also using a binary preference test, we examined whether angelfish are able to discriminate between shoals of small numbers of conspecifics, and if so whether their performance reveals a comparable underlying mechanism to that proposed for discrimination of small quantities in human and non-human animals, namely the possible precursor of the ability to count. Our results demonstrate that fish reliably chose 4 versus 1, 3 versus 1, 2 versus 1 and 3 versus 2 individuals, but were at chance performance level when having to choose between 4 versus 3, 5 versus 4 and 6 versus 5. Findings also reveal that the density of the fish in the stimulus shoals did not significantly affect the performance of experimental angelfish. These results are compatible with the hypothesis of the existence of an object-file mechanism to discriminate small quantities in vertebrates and provide evidence for spontaneous discrimination of up to three elements in angelfish, a similar limit to that found in human and non-human animals. The findings add to the growing body of data, suggesting that the mechanisms underlying discrimination between different quantities of items may be shared across different taxa and have an evolutionary ancient origin.     

Experimental setting affects the performance of guppies in a numerical discrimination task

A recent study found that guppies (Poecilia reticulata) can be trained to discriminate 4 versus 5 objects, a numerical discrimination typically achieved only by some mammals and birds. In that study, guppies were required to discriminate between two patches of small objects on the bottom of the tank that they could remove to find a food reward. It is not clear whether this species possesses exceptional numerical accuracy compared with the other ectothermic vertebrates or whether its remarkable performance was due to a specific predisposition to discriminate between differences in the quality of patches while foraging. To disentangle these possibilities, we trained guppies to the same numerical discriminations with a more conventional two-choice discrimination task. Stimuli were sets of dots presented on a computer screen, and the subjects received a food reward upon approaching the set with the larger numerosity. Though the cognitive problem was identical in the two experiments, the change in the experimental setting led to a much poorer performance as most fish failed even the 2 versus 3 discrimination. In four additional experiments, we varied the duration of the decision time, the type of stimuli, the length of training, and whether correction was allowed in order to identify the factors responsible for the difference. None of these parameters succeeded in increasing the performance to the level of the previous study, although the group trained with three-dimensional stimuli learned the easiest numerical task. We suggest that the different results with the two experimental settings might be due to constraints on learning and that guppies might be prepared to accurately estimate patch quality during foraging but not to learn an abstract stimulus–reward association.     

Origins of number sense. Large-number discrimination in human infants

Four experiments investigated infants' sensitivity to large, approximate numerosities in auditory sequences. Prior studies provided evidence that 6-month-old infants discriminate large numerosities that differ by a ratio of 2.0, but not 1.5, when presented with arrays of visual forms in which many continuous variables are controlled. The present studies used a head-turn preference procedure to test for infants' numerosity discrimination with auditory sequences designed to control for element duration, sequence duration, interelement interval, and amount of acoustic energy. Six-month-old infants discriminated 16 from 8 sounds but failed to discriminate 12 from 8 sounds, providing evidence that the same 2.0 ratio limits numerosity discrimination in auditory-temporal sequences and visual-spatial arrays. Nine-month-old infants, in contrast, successfully discriminated 12 from 8 sounds, but not 10 from 8 sounds, providing evidence that numerosity discrimination increases in precision over development, prior to the emergence of language or symbolic counting.     

大小数量加工机制分离现象的ERP研究

大量行为研究表明,个体对4以上非符号数量的加工存在加工机制分离现象,但是相关的神经机制研究还很少。本研究选取21名大学生被试,结合ERP技术和数量判断任务,以数量比率效应为指标,比较4以上不同大小数量加工的神经反应模式。结果显示,大小数量加工条件下数量比率效应对P2p成分幅值的调节模式存在显著差异。这支持了小数量加工基于数量属性,大数量加工基于非数量属性的观点。     

Quantity discrimination in angelfish (<Emphasis Type="Italic">Pterophyllum scalare</Emphasis>) is maintained after a 30-s retention interval in the large but not in the small number range

The ability to discriminate between sets that differ in the number of elements can be useful in different contexts and may have survival and fitness consequences. As such, numerical/quantity discrimination has been demonstrated in a diversity of animal species. In the laboratory, this ability has been analyzed, for example, using binary choice tests. Furthermore, when the different number of items first presented to the subjects are subsequently obscured, i.e., are not visible at the moment of making a choice, the task requires memory for the size of the sets. In previous work, angelfish (Pterophyllum scalare) have been found to be able to discriminate shoals differing in the number of shoal members both in the small (less than 4) and the large (4 or more) number range, and they were able to perform well even when a short memory retention interval (2–15 s) was imposed. In the current study, we increased the retention interval to 30 s during which the shoals to choose between were obscured, and investigated whether angelfish could show preference for the larger shoal they saw before this interval. Subjects were faced with a discrimination between numerically small shoals (≤4 fish) and also between numerically large (≥4 fish) shoals of conspecifics. We found angelfish not to be able to remember the location of larger versus smaller shoals in the small number range, but to exhibit significant memory for the larger shoal in the large number range as long as the ratio between these shoals was at least 2:1. These results, together with prior findings, suggest the existence of two separate quantity estimation systems, the object file system for small number of items that does not work with the longer retention interval and the analogue magnitude system for larger number of items that does.     

Quantity discrimination in female mosquitofish

The ability in animals to count and represent different numbers of objects has received a great deal of attention in the past few decades. Cumulative evidence from comparative studies on number discriminations report obvious analogies among human babies, non-human primates and birds and are consistent with the hypothesis of two distinct and widespread mechanisms, one for counting small numbers (<4) precisely, and one for quantifying large numbers approximately. We investigated the ability to discriminate among different numerosities, in a distantly related species, the mosquitofish, by using the spontaneous choice of a gravid female to join large groups of females as protection from a sexually harassing male. In one experiment, we found that females were able to discriminate between two shoals with a 1:2 numerosity ratio (2 vs. 4, 4 vs. 8 and 8 vs. 16 fish) but failed to discriminate a 2:3 ratio (8 vs. 12 fish). In the second experiment, we studied the ability to discriminate between shoals that differed by one element; females were able to select the larger shoal when the paired numbers were 2 vs. 3 or 3 vs. 4 but not 4 vs. 5 or 5 vs. 6. Our study indicates that numerical abilities in fish are comparable with those of other non-verbal creatures studied; results are in agreement with the hypothesis of the existence of two distinct systems for quantity discrimination in vertebrates.     

Infants’ auditory enumeration: Evidence for analog magnitudes in the small number range

Vigorous debate surrounds the issue of whether infants use different representational mechanisms to discriminate small and large numbers. We report evidence for ratio-dependent performance in infants’ discrimination of small numbers of auditory events, suggesting that infants can use analog magnitudes to represent small values, at least in the auditory domain. Seven-month-old infants in the present study reliably discriminated two from four tones (a 1:2 ratio) in Experiment 1, when melodic and continuous temporal properties of the sequences were controlled, but failed to discriminate two from three tones (a 2:3 ratio) under the same conditions in Experiment 2. A third experiment ruled out the possibility that infants in Experiment 1 were responding to greater melodic variety in the four-tone sequences. The discrimination function obtained here is the same as that found for infants’ discrimination of large numbers of visual and auditory items at a similar age, as well as for that obtained for similar-aged infants’ duration discriminations, and thus adds to a growing body of evidence suggesting that human infants may share with adults and nonhuman animals a mechanism for representing quantities as “noisy” mental magnitudes.     

Discrimination of small quantities by fish (redtail splitfin, Xenotoca eiseni)

Discrimination of quantity has been argued to rely on two non-verbal representational systems: an object file system (OFS) for representing small values (≤3–4) and an analog magnitude system (AMS) for representing large magnitudes (>4). Infants’ ability to discriminate 1 versus 2, 1 versus 3, 2 versus 3, but not 1 versus 4 or 2 versus 4 seems to prove the independence of such systems. Here, we show that redtail splitfin fish (Xenotoca eiseni) performed relative quantity estimations preferring to approach the location previously occupied by the larger in number between two groups of conspecifics (no longer visible at test) in sets of 1 versus 2 and 2 versus 3 items, but failed at 3 versus 4 items, thus showing the same set-size limit as infants for discrimination of small quantities. However, when tested with quantities that spanned the boundary of the two systems, that is, 1 versus 4 and 2 versus 4, fish succeeded. These results thus point to either the use of continuous physical variables and/or the use of the AMS also for small numerousness in fish in this task.     

Enumeration of small and large numerosities: the effect of element visibility

Precise enumeration is associated with small numerosities within the subitizing range (<4 items), while approximate enumeration is associated with large numerosities (>4 items). To date, there is still debate on whether a single continuous process or dual mutually exclusive processes mediate enumeration of small and large numerosities. Here, we evaluated a compromise between these two notions: that the precise representation of number is limited to small numerosities, but that the approximate representation of numerosity spans across both small and large numerosities. We assessed the independence of precise and approximate enumeration by looking at how luminance contrast affected enumeration of elements that differ by ones (1-8) or by tens (10-80). We found that enumeration functions of ones and tens have different characteristics, which is consistent with the presence of two number systems. Subitizing was preserved for small numerosities. However, simply decreasing element visibility changed the variability signatures of small numerosities to match those of large numerosities. Together, our results suggest that small numerosities are mediated by both precise and approximate representations of numerosity.     

Enumeration of small and large numerosities: The effect of element visibility

Precise enumeration is associated with small numerosities within the subitizing range (<4 items), while approximate enumeration is associated with large numerosities (>4 items). To date, there is still debate on whether a single continuous process or dual mutually exclusive processes mediate enumeration of small and large numerosities. Here, we evaluated a compromise between these two notions: that the precise representation of number is limited to small numerosities, but that the approximate representation of numerosity spans across both small and large numerosities. We assessed the independence of precise and approximate enumeration by looking at how luminance contrast affected enumeration of elements that differ by ones (1–8) or by tens (10–80). We found that enumeration functions of ones and tens have different characteristics, which is consistent with the presence of two number systems. Subitizing was preserved for small numerosities. However, simply decreasing element visibility changed the variability signatures of small numerosities to match those of large numerosities. Together, our results suggest that small numerosities are mediated by both precise and approximate representations of numerosity.     

Perception of missing fundamental by a species of songbird (Sturnus vulgaris)

Two experiments investigated whether a species of songbird perceives missing fundamentals in sounds containing complex frequencies. In Experiment 1, European starlings were trained to discriminate between two sinusoids. This discrimination persisted when the sinusoids were replaced with waveforms composed solely of four consecutive higher harmonics of the training frequencies. In Experiment 2, starlings trained to discriminate between two complex frequencies consisting of sets of higher harmonics transferred the discrimination to the sinusoidal fundamentals. The results demonstrate that starlings can perceive harmonic or periodic structure, and show that a species of songbird can use harmonic structure to gain information about its auditory environment. The findings, together with those obtained from fish and mammals, suggest that periodicity pitch perception may be a general process in vertebrate hearing.