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1.  Lateralized mechanisms for encoding of object. Behavioral evidence from an animal model: the domestic chick (Gallus gallus) 
In our previous research we reported a leftward-asymmetry in domestic chicks required to identify a target element, on the basis of its ordinal position, in a series of identical elements. Here we re-coded behavioral data collected in previous studies from chicks tested in a task involving a different kind of numerical ability, to study lateralization in dealing with an arithmetic task. Chicks were reared with a set of identical objects representing artificial social companions. On day 4, chicks underwent a free-choice test in which two sets, each composed of a different number of identical objects (5 vs.10 or 6 vs. 9, Experiment 1), were hidden behind two opaque screens placed in front of the chick, one on the left and one on the right side. Objects disappeared, one by one, behind either screen, so that, for example, one screen occluded 5 objects and the other 10 objects. The left-right position of the larger set was counterbalanced between trials. Results show that chicks, in the attempt to rejoin the set with the higher number of social companions, performed better when this was located to the right. However, when the number of elements in the two sets was identical (2 vs. 2, in Experiment 2) and they differed only in the coloration of the objects, this bias was not observed, suggesting a predisposition to map the numerical magnitude from left to right. Future studies should be devoted to the direct investigation of this phenomenon, possibly employing an identical number of mono-chromatic imprinting stimuli in both conditions involving a numerical discrimination and conditions not involving any numerosity difference.
doi:10.3389/fpsyg.2014.00150
PMCID: PMC3932408  PMID: 24605106
number cognition; lateralization; counting; number sense; arithmetic; addition; subtraction; domestic chick
2.  Numerical Abstraction in Young Domestic Chicks (Gallus gallus) 
PLoS ONE  2013;8(6):e65262.
In a variety of circumstances animals can represent numerical values per se, although it is unclear how salient numbers are relative to non-numerical properties. The question is then: are numbers intrinsically distinguished or are they processed as a last resort only when no other properties differentiate stimuli? The last resort hypothesis is supported by findings pertaining to animal studies characterized by extensive training procedures. Animals may, nevertheless, spontaneously and routinely discriminate numerical attributes in their natural habitat, but data available on spontaneous numerical competence usually emerge from studies not disentangling numerical from quantitative cues. In the study being outlined here, we tested animals' discrimination of a large number of elements utilizing a paradigm that did not require any training procedures. During rearing, newborn chicks were presented with two stimuli, each characterized by a different number of heterogeneous (for colour, size and shape) elements and food was found in proximity of one of the two stimuli. At testing 3 day-old chicks were presented with stimuli depicting novel elements (for colour, size and shape) representing either the numerosity associated or not associated with food. The chicks approached the number associated with food in the 5vs.10 and 10vs.20 comparisons both when quantitative cues were unavailable (stimuli were of random sizes) or being controlled. The findings emerging from the study support the hypothesis that numbers are salient information promptly processed even by very young animals.
doi:10.1371/journal.pone.0065262
PMCID: PMC3679104  PMID: 23776457
3.  Spontaneous discrimination of possible and impossible objects by newly hatched chicks 
Biology Letters  2011;7(5):654-657.
Four-month-old infants can integrate local cues provided by two-dimensional pictures and interpret global inconsistencies in structural information to discriminate between possible and impossible objects. This leaves unanswered the issue of the relative contribution of maturation of biologically predisposed mechanisms and of experience with real objects, to the development of this capability. Here we show that, after exposure to objects in which junctions providing cues to global structure were occluded, day-old chicks selectively approach the two-dimensional image that depicted the possible rather than the impossible version of a three-dimensional object, after restoration of the junctions. Even more impressively, completely naive newly hatched chicks showed spontaneous preferences towards approaching two-dimensional depictions of structurally possible rather than impossible objects. These findings suggest that the vertebrate brain can be biologically predisposed towards approaching a two-dimensional image representing a view of a structurally possible three-dimensional object.
doi:10.1098/rsbl.2011.0051
PMCID: PMC3169041  PMID: 21429912
impossible objects; early predispositions; object perception; domestic chick
4.  Summation of Large Numerousness by Newborn Chicks 
Newly hatched domestic chicks, reared with identical objects, when presented with sets of 3 vs. 2 objects disappearing one-by-one behind separate screens, spontaneously inspected the screen occluding the larger set; even when the continuous variables (area or perimeter) were controlled for (Rugani et al., 2009). Here, using a similar paradigm, we investigated the ability of chicks to perform addition on larger sets of objects. Chicks imprinted on five identical objects, were presented at test with 6 vs. 9 objects which disappeared one-by-one (Exp. 1). In Exp. 2, the same overall number of objects (15) was used, but employing an increased ratio, i.e., 5 vs. 10. In both experiments, when continuous variables were not made equal, chicks spontaneously inspected the screen occluding the larger set. However, when the size of the objects was adjusted so as to make the total surface area or perimeter equal for the two sets, chicks did not exhibit any preference. Lack of choice in the control conditions could be due to a combination of preferences; to rejoin the larger numerousness as well as the bigger objects (Rugani et al., 2010a). In Exp. 3, chicks were familiarized, during imprinting, with objects of various dimensions, in an attempt to reduce or suppress their tendency to approach objects larger than the familiar ones. Again chicks failed to choose at test between 5 vs. 10 objects when continuous variables were made equal. Results showed that chicks, after a one-by-one presentation of a large number of objects, rejoined the larger set. In order to choose the larger set, chicks estimated the objects in the two sets and then compared the outcomes. However, differently to what has been described for small numerousness, chicks succeeded only if non-numerical cues as well as numerical cues were available. This study suggests that continuous variables are computed by chicks for sets of objects that are not present at the same time and that are no longer visible at the time of choice.
doi:10.3389/fpsyg.2011.00179
PMCID: PMC3171108  PMID: 21941514
number cognition; counting; number sense; arithmetic; domestic chick; human infant; large numeracy; continuous variables
5.  Empty Sets as Part of the Numerical Continuum: Conceptual Precursors to the Zero Concept in Rhesus Monkeys 
The goal of the current research was to explore whether monkeys possess conceptual precursors necessary for understanding zero. We trained rhesus monkeys on a nonsymbolic numerical matching-to-sample task, and on a numerical ordering task. We then introduced nondifferentially reinforced trials that contained empty sets to determine whether monkeys would treat empty sets as numerical values. All monkeys successfully matched and ordered the empty sets without any training. Accuracy showed distance effects, indicating that they treated empty sets as values on a numerical continuum.
doi:10.1037/a0015231
PMCID: PMC2918401  PMID: 19397383
zero; empty sets; numerical cognition; distance effects; numerical continuum
6.  Arithmetic in newborn chicks 
Newly hatched domestic chicks were reared with five identical objects. On days 3 or 4, chicks underwent free-choice tests in which sets of three and two of the five original objects disappeared (either simultaneously or one by one), each behind one of two opaque identical screens. Chicks spontaneously inspected the screen occluding the larger set (experiment 1). Results were confirmed under conditions controlling for continuous variables (total surface area or contour length; experiment 2). In the third experiment, after the initial disappearance of the two sets (first event, FE), some of the objects were visibly transferred, one by one, from one screen to the other (second event, SE). Thus, computation of a series of subsequent additions or subtractions of elements that appeared and disappeared, one by one, was needed in order to perform the task successfully. Chicks spontaneously chose the screen, hiding the larger number of elements at the end of the SE, irrespective of the directional cues provided by the initial (FE) and final (SE) displacements. Results suggest impressive proto-arithmetic capacities in the young and relatively inexperienced chicks of this precocial species.
doi:10.1098/rspb.2009.0044
PMCID: PMC2690459  PMID: 19364746
number cognition; counting; number sense; arithmetic; addition; subtraction
7.  Lateralization of social cognition in the domestic chicken (Gallus gallus) 
In this paper, we report on the ongoing work in our laboratories on the effect of lateralization produced by light exposure in the egg on social cognition in the domestic chick (Gallus gallus). The domestic chick possesses a lateralized visual system. This has effects on the chick's perception towards and interaction with its environment. This includes its ability to live successfully within a social group. We show that there is a tendency for right brain hemisphere dominance when performing social cognitive actions. As such, chicks show a left hemispatial bias for approaching a signalled target object, tend to perceive gaze and faces of human-like masks more effectively when using their left eye, are able to inhibit a pecking response more effectively when viewing a neighbour tasting a bitter substance with their left eye, and are better able to perform a transitive inference task when exposed to light in the egg and when forced to use their left eye only compared to dark-hatched or right eye chicks. Some of these effects were sex specific, with male chicks tending to show an increased effect of lateralization on their behaviours. These data are discussed in terms of overall social cognition in group living.
doi:10.1098/rstb.2008.0229
PMCID: PMC2666078  PMID: 19064355
chick; Gallus gallus; lateralization; social cognition; brain hemisphere
8.  Is it only humans that count from left to right? 
Biology Letters  2010;6(3):290-292.
We report that adult nutcrackers (Nucifraga columbiana) and newborn domestic chicks (Gallus gallus) show a leftward bias when required to locate an object in a series of identical ones on the basis of its ordinal position. Birds were trained to peck at either the fourth or sixth element in a series of 16 identical and aligned positions. These were placed in front of the bird, sagittally with respect to its starting position. When, at test, the series was rotated by 90┬░ lying frontoparallel to the bird's starting position, both species showed a bias for identifying selectively the correct position from the left but not from the right end. The similarity with the well-known phenomenon of the left-to-right spatially oriented number line in humans is considered.
doi:10.1098/rsbl.2009.0960
PMCID: PMC2880063  PMID: 20071393
avian brain; mental number line; domestic chick; Clark's nutcracker

Results 1-8 (8)