Sensitivity to the role of an animated agent from observed interactions in newborn chicks (Gallus gallus).

Few month old human infants are able to detect the social roles of artificial agents and consistently choose the object behaving as 'approacher' rather than 'repulser'. This preference has been considered evidence of a pre-linguistic and pre-cultural origin of the social mind. Similar preferences have not been described in other species, though comparative data could help clarify the nature of this phenomenon and its evolutionary origin. In this study, we investigated sensitivity to the social role of an artificial agent in domestic chicks. Birds offer an excellent model to study the evolutionary roots of cognitive abilities, since they separated from mammals over 300 Ma. Moreover, the investigation of newly hatched chicks allows control for previous experience. After being exposed to computer-presented animations depicting an interaction among two agents, chicks underwent a free choice test among those same objects. While no initial evidence of a clear preference emerged from the planned analysis, chicks in the experimental condition showed a preference for the 'approacher' when controlling for side bias, mirroring human infants behaviour. This suggests the existence of an early ability to discriminate agents from their interactions, independent from any social experience.

Space-luminance crossmodal correspondences in domestic chicks.

Crossmodal correspondences are spontaneous associations of non-redundant information across different modalities. Infants and some non-human animals (i.e., chimpanzees, rhesus monkeys, and dogs) showed crossmodal correspondences like adult humans, suggesting a shared origin (at least among mammals) of such a phenomenon. Here we investigate visual-spatial crossmodal correspondences in a precocial avian species, i.e., the domestic chicken. Three-day-old chicks (n = 40) were presented with two (one in the left and one in the right hemispace of an arena) identical panels, either dyed black (low luminance) or white (high luminance). Chicks could circumnavigate either panel to obtain a food reward. Akin to humans, they preferentially chose the left side when presented with black panels and the right side when presented with white panels. The control group (n = 39), tested with grey panels, showed no spatial preference. In light of our results, we discuss crossmodal correspondences in terms of an early available mechanism widespread across different species.

Spontaneous preference for visual cues of animacy in naïve domestic chicks: The case of speed changes.

Animacy perception arises in human adults from motion cues implying an internal energy source to the moving object. The internal energy of the object is often represented by a change in speed. The same features cause preferential attention in infants. We investigated whether speed changes affecting adults' animacy ratings elicit spontaneous social preferences in visually-naïve chicks. Human observers evaluated the similarity between the movement of a red blob stimulus and that of a living creature. The stimulus entered the screen and moved along the azimuth; halfway through its trajectory it could either continue to move at a constant speed or linearly increase in speed. The average speed, the distance covered and the overall motion duration were kept constant. Animacy ratings of humans were higher for accelerating stimuli (Exp. 1). Naïve chicks were then tested for their spontaneous preference for approaching the stimulus moving at a constant speed and trajectory or an identical stimulus, which suddenly accelerated and then decelerated again to the original speed. Chicks showed a significant preference for the 'speed-change stimulus' (Exp. 2). Two additional controls (Exp. 3 and 4) showed that matching the variability of the control 'speed-constant' stimulus to that of the 'speed-change stimulus' did not alter chicks' preference for the latter. Chicks' preference was suppressed by adding two occluders on both displays, positioned along the stimulus trajectory in such a way to occlude the moment of the speed change (Exp. 5). This confirms that, for chicks to show a preference, the moments of speed change need to be visible. Finally, chicks' preference extended to stimuli displaying a direction change, another motion cue eliciting animacy perception in human observers, if the speed- and direction-profile were consistent with each other and resembled what expected for biological entities that invert their motion direction (Exp. 6). Overall, this is the first demonstration of social predispositions for speed changes in any naïve model or non-human animal, indicating the presence of an attentional filter tuned toward one of the general properties of animate creatures. The similarity with human data suggests a phylogenetically old mechanism shared between vertebrates. Finally, the paradigm developed here provides ground for future investigations of the neural basis of these phenomena.

Numerical discrimination by frogs (Bombina orientalis).

Evidence has been reported for quantity discrimination in mammals and birds and, to a lesser extent, fish and amphibians. For the latter species, however, whether quantity discrimination would reflect sensitivity to number or to the continuous physical variables that covary with number is unclear. Here we reported a series of experiments with frogs (Bombina orientalis) tested in free-choice experiments for their preferences for different amounts of preys (Tenebrio molitor larvae) with systematic controls for variables such as surface area, volume, weight, and movement. Frogs showed quantity discrimination in the range of both small (1 vs. 2, 2 vs. 3, but not 3 vs. 4) and large numerousness (3 vs. 6, 4 vs. 8, but not 4 vs. 6), with clear evidence of being able to discriminate numerousness even when continuous physical variables were controlled for in the case of small numerousness (i.e., 1 vs. 2), whereas in the case of large numerousness it remains unclear whether the number or surface areas were dominant. We suggested that task demands are likely to be responsible for the activation of different systems for small and large numerousness and for their relative susceptibility to quantitative stimulus variables.

Perception of the Ebbinghaus illusion in four-day-old domestic chicks (Gallus gallus).

In the Ebbinghaus size illusion, a central circle surrounded by small circles (inducers) appears bigger than an identical one surrounded by large inducers. Previous studies have failed to demonstrate sensitivity to this illusion in pigeons and baboons, leading to the conclusion that avian species (possibly also nonhuman primates) might lack the neural substrate necessary to perceive the Ebbinghaus illusion in a human-like fashion. Such a substrate may have been only recently evolved in the primate lineage. Here, we show that this illusion is perceived by 4-day-old domestic chicks. During rearing, chicks learnt, according to an observational-learning paradigm, to find food in proximity either of a big or of a small circle. Subjects were then tested with Ebbinghaus stimuli: two identical circles, one surrounded by larger and the other by smaller inducers. The percentage of approaches to the perceptually bigger target in animals reinforced on the bigger circle (and vice versa for the other group) was computed. Over four experiments, we demonstrated that chicks are reliably affected by the illusory display. Subjects reinforced on the small target choose the configuration with big inducers, in which the central target appears perceptually smaller; the opposite is true for subjects reinforced on the big target. This result has important implications for the evolutionary history of the neural substrate involved in the perception of the Ebbinghaus illusion.

Long-term memory for a spatial task in young chicks.

Two-day-old chicks, Gallus gallus domesticus, were faced with a spatial task requiring them to make a detour around a U-shaped barrier in order to join a group of conspecifics placed beyond it. Chicks made one detour trial, and were then retested after delays of 30 min, 3 h and 24 h. When retested, the chicks took significantly less time to make the detour, even at 24 h. Chicks that failed to solve the task on the first trial within the time limit (600 s) took as long as naive chicks, when tested again 24 h later, suggesting that long-term memory for the task requires a form of one-trial learning. Since many chicks chose the same direction of detour on both the first trial and the retest, they may have simply shown a stereotyped preferential response. In a further experiment to test this possibility, we used a more complex version of the apparatus: the direction of detour first chosen by the chick was always blocked, and a subsequent detour in the opposite direction was required to complete the task. This time, chicks did not choose the same direction on the first test and when tested 24 h later. However, they proved able to remember the correct direction for at least 24 h if they were allowed to learn the correct path in a series of five trials. Copyright 1999 The Association for the Study of Animal Behaviour.

Detour behaviour, imprinting and visual lateralization in the domestic chick.

Detour behaviour was studied in chicks faced with a vertical-bar barrier behind where an imprinting object (a red ball) was located. Right-eyed chicks took less time to detour the barrier than left-eyed chicks, and binocular chicks showed a bias to detour the barrier on the left side, thus maintaining visual contact with the imprinting object using the lateral field of the right eye, while circling around the barrier. In males, the asymmetries were consistent all along the first two weeks of life, whereas in females they disappeared on days 8 and 11. When tested with a slightly novel version of the original imprinting object (i.e., a ball of a different color), binocular chicks showed a bias to detour the barrier on the right side, thus showing preferential use of the left eye. The same bias occurred when unfamiliar conspecifics were used as goal-objects. Results suggest that cerebral lateralization in birds can directly affect visually-guided motor responses through selective use of the lateral field of vision of the eye contralateral to the hemisphere which has to be put in charge of control of overt behaviour.

Lateral asymmetries due to preferences in eye use during visual discrimination learning in chicks.

Chicks were trained to discriminate between two boxes of the same colour (white) on the basis of their positions using the pecking response. Some chicks were trained to peck at the box on their right side, some at the box on their left side. They were then retrained with two boxes of different colours (one red the other green): in one group of chicks the position of the two boxes was randomly alternated in the various trials (thus making colour a conspicuous but irrelevant cue), in the other it was maintained unchanged. A control group was retrained with two white boxes identical to those used during training. In all of the three groups chicks had to discriminate between the two boxes on the basis of their positions. During training, chicks took less trial and errors to learn when the positive box was placed on their right side and the same occurred during retraining with boxes that maintained a fixed position and during retraining in the control condition. During retraining with position alternation, on the contrary, chicks took less trials and errors to learn when the positive box was placed on their left side. Video recording of the chicks' behaviour while approaching the boxes showed that these lateral asymmetries reflect head and body turning associated to preferences in eye use, likely due to the different specializations of contralateral brain structures. It is argued that position cues engage the right hemisphere, with consequent head turning to the right to allow lateral viewing by the left eye; object-specific cues engage the left hemisphere, with consequent head turning to the left to allow lateral viewing by the right eye.

Lateral asymmetries during responses to novel-coloured objects in the domestic chick: A developmental study.

Chicks were trained to peck on a small coloured (red or green) box for food reinforcement. They were then presented with one box identical to that used during training (familiar) and one of a different colour (unfamiliar). Food reinforcement was delivered for pecking either box, and the right-left position of the two boxes was alternated at random between trials. The number of times chicks pecked at the novel box when it was located on their right or on their left side was recorded. Chicks showed a general tendency to peck more frequently at the novel box when it was located on their right side. The lateral asymmetry was, however, age-dependent. Maximum lateral bias with both red and green novel boxes was observed at around day 4 and day 11 in both sexes, and minimum at around days 7 8 in males and days 8 9 in females. It is suggested that these lateral asymmetries can be accounted for in terms of a head-and-body turning bias associated to preferential use of the lateral field of vision of the left eye, which arises from specializations of the contralateral brain structures (i.e. right hemisphere involvement in response to novelty). The observed changes in lateral asymmetries during development appear, in fact, to fit very well with independent behavioural and pharmacological evidence for biases to left eye use on day 11, and right eye use on day 8.

Perception of partly occluded objects by young chicks.

Completion of partly occluded objects is a ubiquitous phenomenon in human visual perception. It is unclear, however, whether it occurs at all in other species: Studies on visual discrimination learning have revealed that animals usually attend to parts and features of the discriminative stimuli rather than to global object properties. We provide here the first demonstration of recognition of partly occluded objects in a bird species, the domestic chick Gallus gallus, using the naturalistic setting made available by filial imprinting, a process whereby young birds form attachments to their mothers or some artificial substitute. In Experiment 1, newborn chicks were reared singly with a red cardboard triangle, to which they rapidly imprinted and therefore treated as a social partner. On Day 3 of life, the chicks were presented with pairs of objects composed of either isolated fragments or occluded parts of the imprinting stimulus. Chicks consistently chose to associate with complete or with partly occluded versions of the imprinting object rather than with separate fragments of it. Similarly, in Experiment 2, chicks reared with a partly occluded triangle chose to associate with a complete triangle rather than with fragmented one, whereas chicks reared with a fragmented triangle chose to associate with a fragmented triangle and not with a complete one. Newborn chicks thus appear to behave as if they could experience amodal completion.

The development of responses to novel-coloured objects in male and female domestic chicks.

Male and female domestic chicks were trained to peck on a small coloured (red or green) box for food reinforcement. They were then presented with one box identical to that used during training (familiar) and one of a different colour (unfamiliar) and their preferences were assessed in a simultaneous free choice test. The novel colour was green in chicks trained with a red box and red in chicks trained with a green box. Chicks showed marked variations in their preferences for familiarity and novelty as a function of age and sex. In chicks trained with a green box preferences for the familiar colour were stronger in females than in males at all ages of test. A similar sex difference was observed in chicks trained with a red box, except that at around day 9, when males showed temporarily stronger preferences for familiarity than females. When comparing males and females of red-trained chicks to those of green-trained chicks, a shift in colour prefernce, from red to green, was apparent from day 9 in both sexes, though temporarily stronger in females than in males. Levels of preference for the familiar object showed a peak (centred at around day 5/6) and two dips (centred at around day 4/5 and day 10/11) in both sexes irrespective of colour. Results are discussed in relation to current evidence for simultaneous changes in the brain and in the behaviour of young chicks during development.