Debates have arisen as to whether non-human animals actually can learn abstract non-symbolic numerousness or whether they always rely on some continuous physical aspect of the stimuli, covarying with number. Here, we investigated archerfish (Toxotes jaculatrix) non-symbolic numerical discrimination with accurate control for covarying continuous physical stimulus attributes. Archerfish were trained to select one of two groups of black dots (Exp. 1: 3 vs 6 elements; Exp. 2: 2 vs 3 elements); these were controlled for several combinations of physical variables (elements’ size, overall area, overall perimeter, density, and sparsity), ensuring that only numerical information was available. Generalization tests with novel numerical comparisons (2 vs 3, 5 vs 8, and 6 vs 9 in Exp. 1; 3 vs 4, 3 vs 6 in Exp. 2) revealed choice for the largest or smallest numerical group according to the relative number that was rewarded at training. None of the continuous physical variables, including spatial frequency, were affecting archerfish performance. Results provide evidence that archerfish spontaneously use abstract relative numerical information for both small and large numbers when only numerical cues are available.

We found a region of the zebrafish pallium that shows selective activation upon change in the numerosity of visual stimuli. Zebrafish were habituated to sets of small dots that changed in individual size, position, and density, while maintaining their numerousness and overall surface. During dishabituation tests, zebrafish faced a change in number (with the same overall surface), in shape (with the same overall surface and number), or in size (with the same shape and number) of the dots, whereas, in a control group, zebrafish faced the same stimuli as during the habituation. Modulation of the expression of the immediate early genes c-fos and egr-1 and in situ hybridization revealed a selective activation of the caudal part of the dorso-central division of the zebrafish pallium upon change in numerosity. These findings support the existence of an evolutionarily conserved mechanism for approximate magnitude and provide an avenue for understanding its underlying molecular correlates.

Several studies have suggested that vertebrate and invertebrate species may possess a number sense, i.e. an ability to process in a non-symbolic and non-verbal way the numerousness of a set of items. However, this hypothesis has been challenged by the presence of other non-numerical continuous physical variables, which vary along with numerosity (i.e., any change in the number of visual physical elements in a set naturally involves a related change in visual features such as area, density, contour length and convex hull of the stimulus). It is therefore necessary to control and manipulate the continuous physical information when investigating the ability of humans and other animals to perceive numerousness. During decades of research, different methods have been implemented in order to address this issue, which has implications for experiment replicability and inter-species comparisons, since no general standardized procedure is currently being used. Here we present the ‘Generation of Numerical Elements Images Software’ (GeNEsIS) for the creation of non-symbolic numerical arrays in a standardized and user-friendly environment. The main aim of this tool is to provide researchers in the field of numerical cognition a manageable and precise instrument to produce visual numerical arrays controlled for all the continuous variables. Additionally, we implemented the ability to actively guide stimuli presentation during habituation/dishabituation and dual-choice comparison tasks used in human and comparative research.

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.

It is widely acknowledged that vertebrates can discriminate non-symbolic numerosity using an evolutionarily conserved system dubbed Approximate Number System (ANS). Two main approaches have been used to assess behaviourally numerosity in fish: spontaneous choice tests and operant training procedures. In the first, animals spontaneously choose between sets of biologically-relevant stimuli (e.g., conspecifics, food) differing in quantities (smaller or larger). In the second, animals are trained to associate a numerosity with a reward. Although the ability of fish to discriminate numerosity has been widely documented with these methods, the molecular bases of quantities estimation and ANS are largely unknown. Recently, we combined behavioral tasks with molecular biology assays (e.g c-fos and egr1 and other early genes expression) showing that the thalamus and the caudal region of dorso-central part of the telencephalon seem to be activated upon change in numerousness in visual stimuli. In contrast, the retina and the optic tectum mainly responded to changes in continuous magnitude such as stimulus size. We here provide a review and synthesis of these findings.

Fishes navigate through underwater environments with remarkable spatial precision and memory. Freshwater and seawater species make use of several orientation strategies for adaptative behavior that is on par with terrestrial organisms, and research on cognitive mapping and landmark use in fish have shown that relational and associative spatial learning guide goal-directed navigation not only in terrestrial but also in aquatic habitats. In the past thirty years, researchers explored spatial cognition in fishes in relation to the use of environmental geometry, perhaps because of the scientific value to compare them with land-dwelling animals. Geometric navigation involves the encoding of macrostructural characteristics of space, which are based on the Euclidean concepts of “points”, “surfaces”, and “boundaries”. The current review aims to inspect the extant literature on navigation by geometry in fishes, emphasizing both the recruitment of visual/extra-visual strategies and the nature of the behavioral task on orientation performance.