Magnitude shifts spatial attention from left to right in rhesus monkeys as in the human mental number line.

Humans typically represent numbers and quantities along a left-to-right continuum. Early perspectives attributed number-space association to culture; however, recent evidence in newborns and animals challenges this hypothesis. We investigate whether the length of an array of dots influences spatial bias in rhesus macaques. We designed a touch-screen task that required monkeys to remember the location of a target. At test, monkeys maintained high performance with arrays of 2, 4, 6, or 10 dots, regardless of changes in the array's location, spacing, and length. Monkeys remembered better left targets with 2-dot arrays and right targets with 6- or 10-dot arrays. Replacing the 10-dot array with a long bar, yielded more accurate performance with rightward locations, consistent with an underlying left-to-right oriented magnitude code. Our study supports the hypothesis of a spatially oriented mental magnitude line common to humans and animals, countering the idea that this code arises from uniquely human cultural learning.

Hybro chicks outperform Ross308 in a numerical-ordinal task. Cognitive and behavioral comparisons between 2 broiler strains of newborn domestic chicks (Gallus gallus).

Domestic chickens (Gallus gallus) are among those species subject to intensive selection for production. Among the most widely used broiler strains are the Ross308 and the Hybro. From the perspective of animal production, Ross308 were superior to Hybro in weight gain, final body mass, and feed conversion. Intensive selection is thought to also cause behavioral changes and to negatively affect cognitive abilities. Up to date, though, no evidence has been provided on broiler breeds. The aim of this study was to explore cognitive differences among Hybro and Ross308 chickens by assessing their ordinal-numerical abilities. Chicks learned learnt to find a food reward in the 4th container in a series of 10 identical and sagittally aligned containers. We designed a standard training procedure ensuring that all chicks received the same amount of training. The chicks underwent 2 tests: a sagittal and a fronto-parallel one. In the former test, the series was identical to that experienced during training. In the fronto-parallel test, the series was rotated by 90°, thus left-to-right oriented, to assess the capability of transferring the learnt rule with a novel spatial orientation. In the sagittal test, both chicken hybrids selected the 4th item above chance; interestingly the Hybro outperformed the Ross308 chicks. In the fronto-parallel test, both strains selected the 4th left and the 4th right container above chance; nevertheless, the Hybro chicks were more accurate. Our results support the hypothesis that intense selection for production can influence animal cognition and behavior, with implications on animal husbandry and welfare.

An insect brain organizes numbers on a left-to-right mental number line.

The "mental number line" (MNL) is a form of spatial numeric representation that associates small and large numbers with the left and right spaces, respectively. This spatio-numeric organization can be found in adult humans and has been related to cultural factors such as writing and reading habits. Yet, both human newborns and birds order numbers consistently with an MNL, thus raising the question of whether culture is a main explanation for MNL. Here, we explored the numeric sense of honey bees and show that after being trained to associate numbers with a sucrose reward, they order numbers not previously experienced from left to right according to their magnitude. Importantly, the location of a number on that scale varies with the reference number previously trained and does not depend on low-level cues present on numeric stimuli. We provide a series of neural explanations for this effect based on the extensive knowledge accumulated on the neural underpinnings of visual processing in honey bees and conclude that the MNL is a form of numeric representation that is evolutionarily conserved across nervous systems endowed with a sense of number, irrespective of their neural complexity.

Processing Individually Distinctive Schematic-Faces Supports Proto-Arithmetical Counting in the Young Domestic Chicken.

A key signature of small-number processing is the difficulty in discriminating between three and four objects, as reported in infants and animals. Five-day-old chicks overcome this limit if individually distinctive features characterize each object. In this study, we have investigated whether processing individually different face-like objects can also support discrimination between three and four objects. Chicks were reared with seven face-like stimuli and tested in the proto-arithmetic comparison 1 + 1 + 1 vs. 1 + 1 + 1 + 1. Birds reared and tested with all different faces discriminated and approached the larger group (Exp. 1), whereas new birds reared and tested with seven identical copies of one same face failed (Exp. 2). The presence at test of individually different faces allowed discrimination even when chicks were reared with copies of one face (Exp. 3). To clarify the role of the previous experience of at least one specific arrangement of facial features, in Experiment 4, featureless faces were employed during rearing. During testing, chicks were unable to discriminate between three and four individually distinct faces. Results highlight the importance of having experienced at least one "face" in prompting individual processing and proto-arithmetical calculation later during testing. We speculate that mechanisms effective at the non-symbolic level may positively affect numerical performance.

Are Rational Numbers Spontaneous? Natural Numbers Suffice all Processing by the Number Sense.

The presence of preverbal numerical abilities in animals and infants is widely established, but an important discussion remains about which cognitive systems support these abilities. In particular, a great amount of research is dedicated to the approximate number system (ANS) for the elaboration of non-symbolic numbers and their possible types of mental numeric representations. In a recently published article, Clarke and Beck (2021) provide a series of evidence that supports its existence (ANS) and argue that the mental referents of this system are both natural and rational numbers. In the current commentary, we introduce the notion of the "whole-entity bias" that permeates perception and cognition and favors an automatic processing of discrete mental magnitudes while hindering the representations of fractional numbers. Further, we argue that a hierarchical structure of the ANS that represents natural numbers and supports the understanding of proportions through a series of computations is a more ecological theorization. To conclude, we believe that such a view is still compatible with the proposal of a single representational system (the ANS) supporting both number kinds but offers a different perspective on the computational level of explanation.

Children perform better on left than right targets in an ordinal task.

Francis Galton first reported that humans mentally organize numbers from left to right on a mental number line (1880). This spatial-numerical association was long considered to result from writing and reading habits. More recently though, newborns and animals showed a left-to-right oriented spatial numerical association challenging the primary role assigned to culture in determining the link between number and space. Despite growing evidence supporting the intrinsic association between number and space in different species, its adaptive value is still largely unknown. Here we tested for an advantage in identification of left versus right target positions in 3- to 6-year-old children. Children watched as a toy was hidden under one of 10 linearly arranged identical cups and were then asked to help a stuffed animal retrieve the toy. On each trial, the toy was hidden in the 2nd, 3rd, or 4th cup, from the left or right. To prevent children from staring at the target cup, they were asked to pick up the stuffed animal from under their chair after witnessing the hiding of the toy and then to help the stuffed animal find the toy. Older children were more accurate than younger children. Children exhibited a serial position effect, with performance higher for more exterior targets. Remarkably, children also showed a left bias: they remembered the left targets better than the right targets. Only the youngest children were dramatically influenced by the location of the experimenter during search. Additional analyses support the hypothesis that children used a left-to-right oriented searching strategy in this spatial/ordinal task.

Relative numerical middle in rhesus monkeys.

Animals show vast numerical competence in tasks that require both ordinal and cardinal numerical representations, but few studies have addressed whether animals can identify the numerical middle in a sequence. Two rhesus monkeys (Macaca mulatta) learned to select the middle dot in a horizontal sequence of three dots on a touchscreen. When subsequently presented with longer sequences composed of 5, 7 or 9 items, monkeys transferred the middle rule. Accuracy decreased as the length of the sequence increased. In a second test, we presented monkeys with asymmetrical sequences composed of nine items, where the numerical and spatial middle were distinct and both monkeys selected the numerical middle over the spatial middle. Our results demonstrate that rhesus macaques can extract an abstract numerical rule to bisect a discrete set of items.

Approach direction and accuracy, but not response times, show spatial-numerical association in chicks.

Chicks trained to identify a target item in a sagittally-oriented series of identical items show a higher accuracy for the target on the left, rather than that on the right, at test when the series was rotated by 90°. Such bias seems to be due to a right hemispheric dominance in visuospatial tasks. Up to now, the bias was highlighted by looking at accuracy, the measure mostly used in non-human studies to detect spatial numerical association, SNA. In the present study, processing by each hemisphere was assessed by scoring three variables: accuracy, response times and direction of approach. Domestic chicks were tested under monocular vision conditions, as in the avian brain input to each eye is mostly processed by the contralateral hemisphere. Four-day-old chicks learnt to peck at the 4th element in a sagittal series of 10 identical elements. At test, when facing a series oriented fronto-parallel, birds confined their responses to the visible hemifield, with high accuracy for the 4th element. The first element in the series was also highly selected, suggesting an anchoring strategy to start the proto-counting at one end of the series. In the left monocular condition, chicks approached the series starting from the left, and in the right monocular condition, they started from the right. Both hemispheres appear to exploit the same strategy, scanning the series from the most lateral element in the clear hemifield. Remarkably, there was no effect in the response times: equal latency was scored for correct or incorrect and for left vs. right responses. Overall, these data indicate that the measures implying a direction of choice, accuracy and direction of approach, and not velocity, i.e., response times, can highlight SNA in this paradigm. We discuss the relevance of the selected measures to unveil SNA.

Response of male and female domestic chicks to change in the number (quantity) of imprinting objects.

When facing two sets of imprinting objects of different numerousness, domestic chicks prefer to approach the larger one. Given that choice for familiar and novel stimuli in imprinting situations is known to be affected by the sex of the animals, we investigated how male and female domestic chicks divide the time spent in the proximity of a familiar versus an unfamiliar number of objects, and how animals interact (by pecking) with these objects. We confirmed that chicks discriminate among the different numerousnesses, but we also showed that females and males behave differently, depending on the degree of familiarity of the objects. When objects in the testing sets were all familiar, females equally explored both sets and pecked at all objects individually. Males instead selectively approached the familiar numerousness and pecked more at it. When both testing sets comprised familiar as well as novel objects, both males and females approached the larger numerousness of familiar objects. However, chicks directed all their pecks toward the novel object within the set. Differences in the behavior of males and females can be accounted for in terms of sex difference in the motivation to reinstate social contact with the familiar objects and to explore novel ones, likely associated with the ecology and the social structure of the species before domestication.

Individually distinctive features facilitate numerical discrimination of sets of objects in domestic chicks.

Day-old domestic chicks approach the larger of two groups of identical objects, but in a 3 vs 4 comparison, their performance is random. Here we investigated whether adding individually distinctive features to each object would facilitate such discrimination. Chicks reared with 7 objects were presented with the operation 1 + 1 + 1 vs 1 + 1 + 1 + 1. When objects were all identical, chicks performed randomly, as expected (Experiment 1). In the remaining experiments, objects differed from one another due to additional features. Chicks succeeded when those features were differently oriented segments (Experiment 2) but failed when the features were arranged to depict individually different face-like displays (Experiment 3). Discrimination was restored if the face-like stimuli were presented upside-down, disrupting global processing (Experiment 4). Our results support the claim that numerical discrimination in 3 vs 4 comparison benefits from the presence of distinctive features that enhance object individuation due to individual processing. Interestingly, when the distinctive features are arranged into upright face-like displays, the process is susceptible to global over local interference due to configural processing. This study was aimed at assessing whether individual object processing affects numerical discrimination. We hypothesise that in humans similar strategies aimed at improving performance at the non-symbolic level may have positive effects on symbolic mathematical abilities.

Middle identification for rhesus monkeys is influenced by number but not extent.

concept learning provides a fundamental building block for many cognitive functions in humans. Here we address whether rhesus monkeys (Macaca mulatta) can learn the abstract concept of "middle" in a series of objects. First, we trained monkeys to select the middle dot in a horizontal series of three dots presented on a touchscreen. Monkeys maintained a preference to choose the middle dot despite changes in the appearance, location, and spacing of the horizontal series of dots. They maintained high performance when the color, shape and the length of the stimuli were new, indicating that their responses did not depend upon the particular appearance of the array items. Next, we asked whether monkeys would generalize the middle concept to a 7 dot series. Although accuracy decreased when the number of dots was increased, monkeys continued to preferentially select the middle dot. Our results demonstrate that rhesus macaques can learn to use a middle concept for a discrete set of items.

Numerical magnitude, rather than individual bias, explains spatial numerical association in newborn chicks.

We associate small numbers with the left and large numbers with the right side of space. Recent evidence from human newborns and non-human animals has challenged the primary role assigned to culture, in determining this spatial numerical association (SNA). Nevertheless, the effect of individual spatial biases has not been considered in previous research. Here, we tested the effect of numerical magnitude in SNA and we controlled for itablendividual biases. We trained 3-day-old chicks (Gallus gallus) on a given numerical magnitude (5). Then chicks could choose between two identical, left or right, stimuli both representing either 2, 8, or 5 elements. We computed the percentage of Left-sided Choice (LC). Numerical magnitude, but not individual lateral bias, explained LC: LC2 vs. 2>LC5 vs. 5>LC8 vs. 8. These findings suggest that SNA originates from pre-linguistic precursors, and pave the way to the investigation of the neural correlates of the number space association.

Most of the world modern-day languages are written from left to right ­ but what about numbers? As it turns out, the majority of people also represent numbers using a 'mental line', with smaller numbers on the left and larger numbers on the right. Some researchers argue that this phenomenon results from the way humans learn to read and write: in other words, that it is a by-product of culture, rather than an innate property of the brain. Recent evidence suggests that newborn infants, as well as certain species of monkeys and birds, also associate smaller numbers with the left and larger numbers with the right side of space. This raises the possibility that human mental number line may stem from an ability that evolved before language, in a common ancestor of humans and other animals. Yet, critics claim that findings in infants and non-human species result from a failure to account for individual biases in responding. To resolve this controversy, Rugani et al. trained three-day-old domestic chicks to approach a target board sporting five red squares. Chicks were then given the choice to approach two identical boards, which would both show two, five or eight red squares. Rugani et al. showed that when both boards had two red squares, the chicks tended to approach the left-hand board more often than the right. By contrast, when both boards had eight red squares, the birds approached the right-hand board more often than the left. Importantly, no left-right bias was observed when the number of red squared remained unchanged (five). These results also could not be explained by individual chicks favoring the left or right side. Instead, the findings suggest that even newborn animals tend to associate numbers with positions on a mental number line. Additional research is needed to determine the role of experience ­ or culture ­ in shaping this tendency, and future studies should also examine which brain regions support the association between number and space.

Hemispheric specialization in spatial versus ordinal processing in the day-old domestic chick (Gallus gallus).

Different species show an intriguing similarity in representing numerosity in space, starting from left to right. This bias has been attributed to a right hemisphere dominance in processing spatial information. Here, to disentangle the role of each hemisphere in dealing with spatial versus ordinal-numerical information, we tested domestic chicks during monocular versus binocular vision. In the avian brain, the contralateral hemisphere mainly processes the visual input from each eye. Four-day-old chicks learned to peck at the fourth element in a sagittal series of 10 identical elements. At testing, chicks faced a left-to-right-oriented series where the interelement distance was manipulated so that the third element was where the fourth had been at training; this compelled chicks to use either spatial or ordinal cues. Chicks tested binocularly selected both the fourth left and (to a lesser extent) right elements. Chicks tested monocularly chose the third and fourth elements on the seeing side equally. Interhemispheric cooperation resulted in the use of ordinal-numerical information, while each single hemisphere could rely on spatial or ordinal-numerical cue. Both hemispheres can process spatial and ordinal-numerical information, but their interaction results in the supremacy of processing the ordinal-numerical cue.

Use of numerical and spatial information in ordinal counting by zebrafish.

The use of non-symbolic numerical information is widespread throughout the animal kingdom, providing adaptive benefits in several ecological contexts. Here we provide the possible evidence of ordinal numerical skills in zebrafish (Danio rerio). Zebrafish were trained to identify the second exit in a series of five identically-spaced exits along a corridor. When at test the total length of the corridor (Exp. 1) or the distance between exits (Exp. 2) was changed, zebrafish appeared not to use the absolute spatial distance. However, zebrafish relied both on ordinal as well as spatial cues when the number of exits was increased (from 5 to 9) and the inter-exit distance was reduced (Exp. 3), suggesting that they also take into account relative spatial information. These results highlight that zebrafish may provide a useful model organism for the study of the genetic bases of non-symbolic numerical and spatial cognition, and of their interaction.

A mental number line in human newborns.

Humans represent numbers on a mental number line with smaller numbers on the left and larger numbers on the right side. A left-to-right oriented spatial-numerical association, (SNA), has been demonstrated in animals and infants. However, the possibility that SNA is learnt by early exposure to caregivers' directional biases is still open. We conducted two experiments: in Experiment 1, we tested whether SNA is present at birth and in Experiment 2, we studied whether it depends on the relative rather than the absolute magnitude of numerousness. Fifty-five-hour-old newborns, once habituated to a number (12), spontaneously associated a smaller number (4) with the left and a larger number (36) with the right side (Experiment 1). SNA in neonates is not absolute but relative. The same number (12) was associated with the left side rather than the right side whenever the previously experienced number was larger (36) rather than smaller (4) (Experiment 2). Control on continuous physical variables showed that the effect is specific of discrete magnitudes. These results constitute strong evidence that in our species SNA originates from pre-linguistic and biological precursors in the brain.

Numerical Affordance Influences Action Execution: A Kinematic Study of Finger Movement.

Humans represent symbolic numbers as oriented from left to right: the mental number line (MNL). Up to now, scientific studies have mainly investigated the MNL by means of response times. However, the existing knowledge on the MNL can be advantaged by studies on motor patterns while responding to a number. Cognitive representations, in fact, cannot be fully understood without considering their impact on actions. Here we investigated whether a motor response can be influenced by number processing. Participants seated in front of a little soccer goal. On each trial they were visually presented with a numerical (2, 5, 8) or a non-numerical ($) stimulus. They were instructed to kick a small ball with their right index toward a frontal soccer goal as soon as a stimulus appeared on a screen. However, they had to refrain from kicking when number five was presented (no-go signal). Our main finding is that performing a kicking action after observation of the larger digit proved to be more efficient: the trajectory path was shorter and lower on the surface, velocity peak was anticipated. The smaller number, instead, specifically altered the temporal and spatial aspects of trajectories, leading to more prolonged left deviations. This is the first experimental demonstration that the reaching component of a movement is influenced by number magnitude. Since this paradigm does not require any verbal skill and non-symbolic stimuli (array of dots) can be used, it could be fruitfully adopted to evaluate number abilities in children and even preschoolers. Notably, this is a self-motivating and engaging task, which might help children to get involved and to reduce potential arousal connected to institutional paper-and-pencil examinations.

The effect of clustering on perceived quantity in humans (Homo sapiens) and in chicks (Gallus gallus).

Animals can perceive the numerosity of sets of visual elements. Qualitative and quantitative similarities in different species suggest the existence of a shared system (approximate number system). Biases associated with sensory properties are informative about the underlying mechanisms. In humans, regular spacing increases perceived numerosity (regular-random numerosity illusion). This has led to a model that predicts numerosity based on occupancy (a measure that decreases when elements are close together). We used a procedure in which observers selected one of two stimuli and were given feedback with respect to whether the choice was correct. One configuration had 20 elements and the other 40, randomly placed inside a circular region. Participants had to discover the rule based on feedback. Because density and clustering covaried with numerosity, different dimensions could be used. After reaching a criterion, test trials presented two types of configurations with 30 elements. One type had a larger interelement distance than the other (high or low clustering). If observers had adopted a numerosity strategy, they would choose low clustering (if reinforced with 40) and high clustering (if reinforced with 20). A clustering or density strategy predicts the opposite. Human adults used a numerosity strategy. Chicks were tested using a similar procedure. There were two behavioral measures: first approach response and final circumnavigation (walking behind the screen). The prediction based on numerosity was confirmed by the first approach data. For chicks, one clear pattern from both responses was a preference for the configurations with higher clustering. (PsycINFO Database Record