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Characterizing ontogeny of quantity discrimination in zebrafish
Abstract
A sense of non-symbolic numerical magnitudes is widespread in the animal kingdom and has been documented in adult zebrafish. Here, we investigated the ontogeny of this ability using a group size preference (GSP) task in juvenile zebrafish. Fish showed GSP from 21 days post-fertilization and reliably chose the larger group when presented with discriminations of between 1 versus 3, 2 versus 5 and 2 versus 3 conspecifics but not 2 versus 4 conspecifics. When the ratio between the number of conspecifics in each group was maintained at 1 : 2, fish could discriminate between 1 versus 2 individuals and 3 versus 6, but again, not when given a choice between 2 versus 4 individuals. These findings are in agreement with studies in other species, suggesting the systems involved in quantity representation do not operate separately from other cognitive mechanisms. Rather they suggest quantity processing in fishes may be the result of an interplay between attentional, cognitive and memory-related mechanisms as in humans and other animals. Our results emphasize the potential of the use of zebrafish to explore the genetic and neural processes underlying the ontogeny and function of number cognition.
... Zebrafish are physiologically homologous to mammals and possess all major neurotransmitters, hormones, and receptors (Panula et al., 2006;Alsop and Vijayan, 2009). The high degree of protein and genetic homology with humans (Howe et al., 2013), coupled with refined gene-editing tools and behavioral paradigms, make this species a vertebrate system amenable to large-scale forward genetic analyses (Wolman et al., 2011;Kalueff et al., 2013;Sheardown et al., 2022). Transparency of embryos and larvae enables in vivo functional imaging of neural activity and establishes the zebrafish as a powerful optogenetic tool. ...
... Altogether, these results suggest a limited role of maturation and experience on zebrafish's ability to estimate areas that parallel the existence of tectal neurons by selectively responding to item size already during the first week of life (see paragraph below for more details). This high accuracy seems to be in contrast with performance observed in juvenile zebrafish tested in a spontaneous choice test to investigate the ontogeny of numerical competence (Sheardown et al., 2022). In this study, the authors adapted the procedure previously used to investigate shoal choice discrimination in angelfish, in which the stimulus fish were located in individual compartments that limited movement and avoided fish hiding each other. ...
... The consistent failure to select the larger group between 2 and 4 individuals, whether controlled for space occupied or not, suggests that the fish did not only rely on the overall space as 4 fish occupied two times the space. Hence, the authors suggested that zebrafish may use both numerical and non-numerical information to represent quantities (Agrillo et al., 2016;Leibovich et al., 2017;Sheardown et al., 2022) and that attentional constraints and cognitive and working memory mechanisms may orchestrate numerical competence as hypothesized in both humans and other animals (Hyde, 2011). However, these results may also support the hypothesis of distinct quantification systems characterized by domain and task specificity operating largely independently from the others (Feigenson et al., 2004;Miletto Petrazzini et al., 2014). ...
An ability to estimate quantities, such as the number of conspecifics or the size of a predator, has been reported in vertebrates. Fish, in particular zebrafish, may be instrumental in advancing the understanding of magnitude cognition. We review here the behavioral studies that have described the ecological relevance of quantity estimation in fish and the current status of the research aimed at investigating the neurobiological bases of these abilities. By combining behavioral methods with molecular genetics and calcium imaging, the involvement of the retina and the optic tectum has been documented for the estimation of continuous quantities in the larval and adult zebrafish brain, and the contributions of the thalamus and the dorsal-central pallium for discrete magnitude estimation in the adult zebrafish brain. Evidence for basic circuitry can now be complemented and extended to research that make use of transgenic lines to deepen our understanding of quantity cognition at genetic and molecular levels.
... This nding indicates that numerical abilities in amphibians can be underestimated or misinterpreted. Plasticity in this aspect of number sense is already known in other taxa (e.g., in sh, Bisazza et al., 2010;Sheardown et al., 2022). When measuring the number competence of frogs and toads, it will be crucial to consider ontogeny and plasticity. ...
The ability to perceive group size and discriminate the ontogeny of conspecifics would play a crucial role in the grouping behavior of animals. However, the relative importance of numerical quantity and size-assortative preferences in shaping grouping rules remains poorly understood. In this study, I examined the responses of Miyako toad ( Bufo gargarizans miyakonis ) tadpoles to number quantity and size discrimination by choice tests at different ontogenetic stages (small, medium, and large). The results revealed that small-sized tadpoles in early developmental stages significantly preferred larger numbers (4) compared to smaller ones (1). However, this preference was not observed in later developmental stages (medium and large). And interestingly, when there was no group size bias, size discrimination was not observed in tadpoles, irrespective of their ontogeny. These findings suggest that Miyako toad tadpoles possess innate numerical competence but demonstrate ontogeny-dependent utilization of this ability. Understanding the interplay between numerical quantity and size-assortative preferences in grouping behavior will provide esteemed insights into the adaptive value of number sense in vertebrates and sheds light on evolutionary processes.
... However, if the numerical information is excluded and the two stimuli differ only in their continuous property (cumulative area) guppies no longer discriminate between them (Miletto Petrazzini et al. 2014;Piffer et al. 2013). Findings of a recent study of young zebra fish (Danio rerio) are consistent with these results, at least in terms of number discrimination (Sheardown et al. 2022). In contrast with this, tadpoles of the two anuran species tested (Bufotes balearicus and Pelophylax esculentus) seem to have the capacity to discriminate between only small numbers (Balestrieri et al. 2019), suggesting that they rely more on the OFS, whereas fishes might operate on both. ...
Quantitative abilities are well described in many species and in diverse life situations, including in the adult domestic cat. However, such abilities have been much less studied during ontogeny. In the present study we examined spontaneous quantity discrimination by pre-weaning age kittens in two-way food choice experiments. In Experiment 1, 26 kittens performed 12 trials with different ratios between the number of same-size food items. In Experiment 2, 24 other kittens performed eight trials with different ratios between the size of two food items. We found, in general, that the kittens discriminated between the different amounts of food and spontaneously chose the larger one, but that their choice was influenced by the ratio of difference. The kittens in Experiment 1 chose the larger number of same-size food items if the ratio was smaller than 0.4 and in Experiment 2 they chose the larger pieces of food if the ratio between the items was smaller than 0.5. Because the kittens’ choice was not influenced by the absolute number of food items or the numerical difference between them in Experiment 1, it suggests that their cognitive performance relied on an analog magnitude system rather than on an object file system during the quantity discrimination tasks. We discuss our results considering the ecological and social background of cats and compare it with the performance of previously studied species.
... In the aforementioned situations, pure numerical abilities provide reliable information for adaptive decisions. Therefore, it is not difficult to explain their presence in animals that are requested to perform such decisions, including adult and juvenile zebrafish 58,59 . What is puzzling, is the presence of numerical abilities in zebrafish at an early developmental stage when they seem to have reasonably no functional role. ...
An intriguing hypothesis to explain the ubiquity of numerical abilities is that all vertebrates are born with hardwired neuronal networks for processing numbers. To date, only studies on human foetuses have clearly supported this hypothesis. Zebrafish hatch 48–72 h after fertilisation with an embryonic nervous system, providing a unique opportunity for investigating this hypothesis. Here, we demonstrated that zebrafish larvae exposed to vertical bars at birth acquired an attraction for bar stimuli and we developed a numerical discrimination task based on this preference. When tested with a series of discriminations of increasing difficulty (1vs.4, 1vs.3, 1vs.2, and 2vs.4 bars), zebrafish larvae reliably selected the greater numerosity. The preference was significant when stimuli were matched for surface area, luminance, density, and convex hull, thereby suggesting a true capacity to process numerical information. Converging results from two phylogenetically distant species suggests that numerical abilities might be a hallmark feature of vertebrates’ brains.
... However, we see evidence for the preference for the larger shoal both in the case of 4 versus 2 and 6 versus 3 dichotomous trials through trajectory density analysis, where the more dense regions were near the larger shoal. Previous studies on shoaling preference report that zebrafish can discriminate between shoal sizes of 6 versus 3 in dichotomous choices, although many studies indicate a failure to discriminate 4 versus 2 shoal sizes 39,51 .However, both those studies also use a different rectangular three-tank shoal choice set up from ours, and one 39 used a perforated divider through which shoaling decisions were based on olfactory and not just visual cues from the display fish. ...
Several organisms, from slime molds to humans, are known to violate normative principles of economic rationality in decision making. In animals, the neural circuitry underlying behaviors that violate or conform to normative rationality is relatively poorly understood. We investigated whether zebrafish, a model organism with a strong suite of functional neuroimaging and genetic manipulation tools, showed rational behavior with respect to the principle of the Independence of Irrelevant Alternatives (IIA). We examined IIA in social decision making by measuring revealed preferences from spatial trajectories of freely swimming individual zebrafish in an arena where they could view and perform shoaling behavior near conspecific zebrafish in adjacent display tanks. IIA was tested in terms of the invariance of shoaling choices between binary and ternary sets of various display fish group sizes. We provide the first report of evidence for violation of IIA in zebrafish, both in terms of the constant ratio rule and in terms of the principle of regularity. This also is a rare example of violation of rationality in a single attribute dimension, and it opens up a range of possibilities to study the neural basis of context dependent decision making.
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.
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.
The ability to represent, discriminate, and perform arithmetic operations on discrete quantities (numerosities) has been documented in a variety of species of different taxonomic groups, both vertebrates and invertebrates. We do not know, however, to what extent similarity in behavioral data corresponds to basic similarity in underlying neural mechanisms. Here, we review evidence for magnitude representation, both discrete (countable) and continuous, following the sensory input path from primary sensory systems to associative pallial territories in the vertebrate brains. We also speculate on possible underlying mechanisms in invertebrate brains and on the role played by modeling with artificial neural networks. This may provide a general overview on the nervous system involvement in approximating quantity in different animal species, and a general theoretical framework to future comparative studies on the neurobiology of number cognition.
Consciousness shared
Humans have tended to believe that we are the only species to possess certain traits, behaviors, or abilities, especially with regard to cognition. Occasionally, we extend such traits to primates or other mammals—species with which we share fundamental brain similarities. Over time, more and more of these supposed pillars of human exceptionalism have fallen. Nieder et al. now argue that the relationship between consciousness and a standard cerebral cortex is another fallen pillar (see the Perspective by Herculano-Houzel). Specifically, carrion crows show a neuronal response in the palliative end brain during the performance of a task that correlates with their perception of a stimulus. Such activity might be a broad marker for consciousness.
Science , this issue p. 1626 ; see also p. 1567
There is considerable evidence that animals are able to discriminate between quantities. Despite the fact that quantitative skills have been extensively studied in adult individuals, research on their development in early life is restricted to a limited number of species. We, therefore, investigated whether 2-month-old puppies could spontaneously discriminate between different quantities of food items. We used a simultaneous two-choice task in which puppies were presented with three numerical combinations of pieces of food (1 vs. 8, 1 vs. 6 and 1 vs. 4), and they were allowed to select only one option. The subjects chose the larger of the two quantities in the 1 vs. 8 and the 1 vs. 6 combinations but not in the 1 vs. 4 combination. Furthermore, the last quantity the puppies looked at before making their choice and the time spent looking at the larger/smaller amounts of food were predictive of the choices they made. Since adult dogs are capable of discriminating between more difficult numerical contrasts when tested with similar tasks, our findings suggest that the capacity to discriminate between quantities is already present at an early age, but that it is limited to very easy discriminations.
Evidence has shown that a variety of vertebrates, including fish, can discriminate collections of visual items on the basis of their numerousness using an evolutionarily conserved system for approximating numerical magnitude (the so-called Approximate Number System, ANS). Here we combine a habituation/dishabituation behavioural task with molecular biology assays to start investigating the neural bases of the ANS in zebrafish. Separate groups of zebrafish underwent a habituation phase with a set of 3 or 9 small red dots, associated with a food reward. The dots changed in size, position and density from trial to trial but maintained their numerousness, and the overall areas of the stimuli was kept constant. During the subsequent dishabituation test, zebrafish faced a change (i) in number (from 3 to 9 or vice versa with the same overall surface), or (ii) in shape (with the same overall surface and number), or (iii) in size (with the same shape and number). A control group of zebrafish was shown the same stimuli as during the habituation. RT-qPCR revealed that the telencephalon and thalamus were characterized by the most consistent modulation of the expression of the immediate early genes c-fos and egr-1 upon change in numerousness; in contrast, the retina and optic tectum responded mainly to changes in stimulus size.
A large body of literature is available on wound healing in humans. Nonetheless, a standardized ex vivo wound model without disruption of the dermal compartment has not been put forward with compelling justification. Here, we present a novel wound model based on application of negative pressure and its effects for epidermal regeneration and immune cell behaviour. Importantly, the basement membrane remained intact after blister roof removal and keratinocytes were absent in the wounded area. Upon six days of culture, the wound was covered with one to three-cell thick K14+Ki67+ keratinocyte layers, indicating that proliferation and migration were involved in wound closure. After eight to twelve days, a multi-layered epidermis was formed expressing epidermal differentiation markers (K10, filaggrin, DSG-1, CDSN). Investigations about immune cell-specific manners revealed more T cells in the blister roof epidermis compared to normal epidermis. We identified several cell populations in blister roof epidermis and suction blister fluid that are absent in normal epidermis which correlated with their decrease in the dermis, indicating a dermal efflux upon negative pressure. Together, our model recapitulates the main features of epithelial wound regeneration, and can be applied for testing wound healing therapies and investigating underlying mechanisms.
To make appropriate decisions, animals need to accumulate sensory evidence. Simple integrator models can explain many aspects of such behavior, but how the underlying computations are mechanistically implemented in the brain remains poorly understood. Here we approach this problem by adapting the random-dot motion discrimination paradigm, classically used in primate studies, to larval zebrafish. Using their innate optomotor response as a measure of decision making, we find that larval zebrafish accumulate and remember motion evidence over many seconds and that the behavior is in close agreement with a bounded leaky integrator model. Through the use of brain-wide functional imaging, we identify three neuronal clusters in the anterior hindbrain that are well suited to execute the underlying computations. By relating the dynamics within these structures to individual behavioral choices, we propose a biophysically plausible circuit arrangement in which an evidence integrator competes against a dynamic decision threshold to activate a downstream motor command.
For social animals, group size discrimination may play a major role in setting the trade-off between the costs and benefits of membership. Several anuran tadpoles show different degrees of social aggregation when exposed to the risk of predation. Despite the importance of aggregative behaviour as an anti-predatory response, the mechanism underlying tadpole choice of the group to join to has not been sufficiently investigated. To establish whether visual cues provide sufficient information to enable tadpoles to choose between aggregations differing in size, we explored the abilities of the larvae of two anuran species (green toad Bufotes balearicus and edible frog Pelophylax esculentus) to discriminate among four numerical combinations of conspecific tadpoles (1 vs. 4, 3 vs. 4, 4 vs. 6 and 4 vs. 8), either in the presence or absence of predatory cues. Our results suggest that in anuran larvae the capacity to discriminate between quantities is limited to small numbers (1 vs. 4 for B. balearicus and both 1 vs. 4 and 3 vs. 4 for P. esculentus). Predator-exposed toad tadpoles stayed longer close to the larger group, supporting aggregation as a major anti-predator behaviour in bufonids, while frog tadpoles showed a preference for the smaller groups, though in predator-free trials only, probably associated with lower intra-specific competition.
Common strains of wildtype zebrafish (Danio rerio) have unique genomic features including SNPs and CNV, but strain information often goes unreported in the literature. As a result, the confounding effects of interstrain variation makes repetition of studies in zebrafish challenging. Here we analyze hepatic mRNA expression patterns between three common zebrafish strains (AB, Tuebingen (TU), and WIK) using Agilent 4 × 44 K gene expression microarrays to establish baseline mRNA expression across strains and between sexes. We observed wide variation in sex-specific gene expression within AB and WIK strains (141 genes in AB and 67 genes in WIK), but no significant variation between sexes within TU. After partitioning the dataset into male and female subsets, we detected 421 unique mRNA transcripts with statistically significant differential expression; 269 mRNA transcripts varied between males, 212 mRNA transcripts varied between females, and 59 mRNA transcripts varied across the three strains, regardless of sex. It is not surprising that mRNA expression profiles differ between sexes and strains, but it is imperative to characterize the differences. These results highlight the complexity of variation within zebrafish and underscore the value of this model system as a valid representation of normal variation present in other species, including humans.
Zika virus (ZIKV) has recently caused a pandemic disease, and many cases of ZIKV infection in pregnant women resulted in abortion, stillbirth, deaths and congenital defects including microcephaly, which now has been proposed as ZIKV congenital syndrome. This study aimed to investigate the in situ immune response profile and mechanisms of neuronal cell damage in fatal Zika microcephaly cases. Brain tissue samples were collected from 15 cases, including 10 microcephalic ZIKV-positive neonates with fatal outcome and five neonatal control flavivirus-negative neonates that died due to other causes, but with preserved central nervous system (CNS) architecture. In microcephaly cases, the histopathological features of the tissue samples were characterized in three CNS areas (meninges, perivascular space, and parenchyma). The changes found were mainly calcification, necrosis, neuronophagy, gliosis, microglial nodules, and inflammatory infiltration of mononuclear cells. The in situ immune response against ZIKV in the CNS of newborns is complex. Despite the predominant expression of Th2 cytokines, other cytokines such as Th1, Th17, Treg, Th9, and Th22 are involved to a lesser extent, but are still likely to participate in the immunopathogenic mechanisms of neural disease in fatal cases of microcephaly caused by ZIKV.
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.
Rudimentary quantification abilities are found in numerous animal species and in human infants all demonstrating the ability to discriminate between quantities differing in numerical size. An open question is whether individuals rely on different underlying systems to discriminate between large (analogue magnitude system (AMS) for number of items exceeding 3) and small quantities (object-file system (OFS) for number of items below 4), or they use only one system (AMS) for the entire number range. The two-system hypothesis has been supported by finding reduced ability to discriminate between quantities that cross the large-small boundary in several species. Recently, the role of cognitive representation, i.e., memory, in quantity discrimination has also been recognized. Here, we investigated whether angelfish can discriminate quantities across the boundary under two memory conditions. In a binary choice test, single angelfish were allowed to see groups (shoals) of conspecifics of different numerical size on the two sides of their test tank. In Experiment 1, their choice was recorded after a 2-sec retention interval during which shoal size information was unavailable. Angelfish were able to discriminate the larger shoal across the boundary when the shoals differed by a 2:1 or higher ratio, but not when the ratio was lower. In Experiment 2, however, with a 15-sec retention interval, angelfish could only detect a four-fold difference in ratio but failed to detect a three- or a two-fold difference across the boundary. These results suggest that angelfish can remember smaller differences for a short (2 sec) but not for a longer (15 sec) period. Together with previous findings, the current results support the idea that angelfish use two distinct systems for representing quantity, but they may recruit the AMS even for the small number range under some circumstances, e.g., when higher memory demand is imposed by a greater retention interval.
Adult zebrafish are robustly social animals whereas larva is not. We designed an assay to determine at what stage of development zebrafish begin to interact with and prefer other fish. One week old zebrafish do not show significant social preference whereas most 3 weeks old zebrafish strongly prefer to remain in a compartment where they can view conspecifics. However, for some individuals, the presence of conspecifics drives avoidance instead of attraction. Social preference is dependent on vision and requires viewing fish of a similar age/size. In addition, over the same 1-3 weeks period larval zebrafish increasingly tend to coordinate their movements, a simple form of social interaction. Finally, social preference and coupled interactions are differentially modified by an NMDAR antagonist and acute exposure to ethanol, both of which are known to alter social behavior in adult zebrafish.
In this study we investigated how the networks mediating respiratory and locomotor drives to lumbar motoneurons interact and how this interaction is modulated in relation to periodic variations in blood pressure (Mayer waves). Seven decerebrate cats, under neuromuscular blockade, were used to study central respiratory drive potentials (CRDPs, usually enhanced by added CO2) and spontaneously occurring locomotor drive potentials (LDPs) in hindlimb motoneurons, together with hindlimb and phrenic nerve discharges. In four of the cats both drives and their voltage-dependent amplification were absent or modest, but in the other three, one or other of these drives was common and the voltage-dependent amplification was frequently strong. Moreover, in these three cats the blood pressure showed marked periodic variation (Mayer waves), with a slow rate (periods 9-104 s, mean 39 ± 17 SD). Profound modulation, synchronized with the Mayer waves was seen in the occurrence and/or in the amplification of the CRDPs or LDPs. In one animal, where CRDPs were present in most cells and the amplification was strong, the CRDP consistently triggered sustained plateaux at one phase of the Mayer wave cycle. In the other two animals, LDPs were common, and the occurrence of the locomotor drive was gated by the Mayer wave cycle, sometimes in alternation with the respiratory drive. Other interactions between the two drives involved respiration providing leading events, including co-activation of flexors and extensors during post-inspiration or a locomotor drive gated or sometimes entrained by respiration. We conclude that the respiratory drive in hindlimb motoneurons is transmitted via elements of the locomotor central pattern generator. The rapid modulation related to Mayer waves suggests the existence of a more direct and specific descending modulatory control than has previously been demonstrated.
Due to their well-characterized neural development and high genetic homology to mammals, zebrafish (Danio rerio) have emerged as a powerful model organism in the field of biological psychiatry. Here, we discuss the molecular psychiatry of zebrafish, and its implications for translational neuroscience research and modeling central nervous system (CNS) disorders. In particular, we outline recent genetic and technological developments allowing for in vivo examinations, high-throughput screening and whole-brain analyses in larval and adult zebrafish. We also summarize the application of these molecular techniques to the understanding of neuropsychiatric disease, outlining the potential of zebrafish for modeling complex brain disorders, including attention-deficit/hyperactivity disorder (ADHD), aggression, post-traumatic stress and substance abuse. Critically evaluating the advantages and limitations of larval and adult fish tests, we suggest that zebrafish models become a rapidly emerging new field in modern molecular psychiatry research.Molecular Psychiatry advance online publication, 28 October 2014; doi:10.1038/mp.2014.128.
Basic intellectual abilities of quantity and numerosity estimation have been detected across animal species. Such abilities are referred to as ‘number sense’. For human species, individual differences in number sense are detectable early in life, persist in later development, and relate to general intelligence. The origins of these individual differences are unknown. To address this question, we conducted the first large-scale genetically sensitive investigation of number sense, assessing numerosity discrimination abilities in 837 pairs of monozygotic and 1422 pairs of dizygotic 16-year-old twin pairs. Univariate genetic analysis of the twin data revealed that number sense is modestly heritable (32%), with individual differences being largely explained by non-shared environmental influences (68%) and no contribution from shared environmental factors. Sex-Limitation model fitting revealed no differences between males and females in the etiology of individual differences in number sense abilities. We also carried out Genome-wide Complex Trait Analysis (GCTA) that estimates the population variance explained by additive effects of DNA differences among unrelated individuals. For 1118 unrelated individuals in our sample with genotyping information on 1.7 million DNA markers, GCTA estimated zero heritability for number sense, unlike other cognitive abilities in the same twin study where the GCTA heritability estimates were about 25%. The low heritability of number sense, observed in this study, is consistent with the directional selection explanation whereby additive genetic variance for evolutionary important traits is reduced.
Human adults and nonhuman primates share a subset of nonverbal numerical skills that are considered the evolutionary foundation of more complex numerical reasoning. Intriguing experiments have shown that 10- to 12-month-old infants are able to distinguish between large (8 vs. 12) and small (1 vs. 2, 1 vs. 3, 2 vs. 3) sets of objects but seem incapable of comparing quantities that fall in the middle area between large and small numerosities, such as 1 versus 4. This finding suggests that there are two separate nonverbal numerical systems. Other researchers argue that there is continuity in the representation of numbers. Experimental evidence demonstrating that newborn chicks are able to process addition and subtraction such as (4-1) versus (1 + 1) lends support to the latter hypothesis. Here, using an experimental paradigm to test numerical discrimination, we demonstrated that newborn chicks are able to distinguish between some numerical comparisons, such as 2 vs. 3, 2 vs. 8, 6 vs. 9, 8 vs. 14, 4 vs. 6, and 4 vs. 8. These findings support the hypothesis that a single system processes both small and large numerosities. The results of these experiments demonstrate that small and large numbers can be discriminated via "analogue magnitude" system (AMS). Those data can be accounted for in terms of a select mechanism prompting the functioning of either system and, therefore, a different processing of the stimuli. When the modality of presentation of the stimuli focuses the attention on the whole collection, the elaboration would be carried out by the AMS. (PsycINFO Database Record (c) 2013 APA, all rights reserved).
Subitizing, the fast and accurate enumeration of up to about 3 or 4 objects, has often been thought to be dependent on limited-capacity preattentive mechanisms. We used an attentional blink paradigm to investigate the extent to which subitizing requires attentional resources. On each trial, subjects identified a target letter in an RSVP stream and then enumerated dots presented in the stream that were either simultaneous with the target letter or followed it by up to 400 ms. For numerosities from 2 to 9, evidence of an attentional blink was observed; only enumeration of 0 or 1 elements was independent of lag. Thus, even enumeration of 2–3 objects, which is within the traditional subitizing range, appears to require attentional resources. The relation of this work to studies on the attentional requirements of detecting a unique item among distractors, a supposedly preattentive discrimination, is briefly discussed.
Zebrafish have become a popular organism for the study of vertebrate gene function. The virtually transparent embryos of this species, and the ability to accelerate genetic studies by gene knockdown or overexpression, have led to the widespread use of zebrafish in the detailed investigation of vertebrate gene function and increasingly, the study of human genetic disease. However, for effective modelling of human genetic disease it is important to understand the extent to which zebrafish genes and gene structures are related to orthologous human genes. To examine this, we generated a high-quality sequence assembly of the zebrafish genome, made up of an overlapping set of completely sequenced large-insert clones that were ordered and oriented using a high-resolution high-density meiotic map. Detailed automatic and manual annotation provides evidence of more than 26,000 protein-coding genes, the largest gene set of any vertebrate so far sequenced. Comparison to the human reference genome shows that approximately 70% of human genes have at least one obvious zebrafish orthologue. In addition, the high quality of this genome assembly provides a clearer understanding of key genomic features such as a unique repeat content, a scarcity of pseudogenes, an enrichment of zebrafish-specific genes on chromosome 4 and chromosomal regions that influence sex determination.
Whereas cognitive impairment is a common symptom in multiple brain disorders, predictive and high-throughput animal models of cognition and behavior are becoming increasingly important in the field of translational neuroscience research. In particular, reliable models of the cognitive deficits characteristic of numerous neurobehavioral disorders such as Alzheimer's disease and schizophrenia have become a significant focus of investigation. While rodents have traditionally been used to study cognitive phenotypes, zebrafish () are gaining popularity as an excellent model to complement current translational neuroscience research. Here we discuss recent advances in pharmacological and genetic approaches using zebrafish models to study cognitive impairments and to discover novel cognitive enhancers and neuroprotective mechanisms.
Human adults master sophisticated, abstract numerical calculations that are mostly based on symbolic language and thus inimitably human. Humans may nonetheless share a subset of non-verbal numerical skills, available soon after birth and considered the evolutionary foundation of more complex numerical reasoning, with other animals. These skills are thought to be based on the two systems: the object file system which processes small values (<3) and the analogue magnitude system which processes large magnitudes (>4). Infants' ability to discriminate 1 vs. 2, 1 vs. 3, 2 vs. 3, but not 1 vs. 4, seems to indicate that the two systems are independent, implying that the conception of a continuous number processing system is based on precursors that appear to be interrupted at or about the number four. The findings from the study being presented here indicating that chicks are able to make a series of discriminations regarding that borderline number (1 vs. 4, 1 vs. 5, 2 vs. 4) support the hypothesis that there is continuity in the number system which processes both small and large numerousness.
Human infants and non-human animals can discriminate the larger of two sets of discrete items. This quantity discrimination may be based upon the number of items, or upon non-numerical variables of the sets that co-vary with number. We have demonstrated that angelfish select the larger of two shoals of conspecifics without using inter-fish distance or space occupied by the stimuli as cues. However, density appeared to influence the choice between large shoals. Here, we examine the role of another non-numerical cue, swimming activity of the stimulus fish, in quantity discrimination by angelfish. To control this variable, we varied the water temperature of the stimulus aquaria or restricted the space occupied by each fish in the stimulus shoals. We used the previously successfully discriminated contrasts consisting of large (10 vs. 5) and small (3 vs. 2) shoals. We also studied whether more active or less active shoals are preferred in case of equally sized shoals (10 vs. 10, 5 vs. 5, and 3 vs. 3). When differences in stimulus fish activity were minimized by temperature manipulation we found angelfish to prefer the larger shoal in the 3 vs. 2 comparison, but not in the 10 vs. 5 comparison. When activity was controlled by space restriction, angelfish preferred the larger shoal in both numerical contrasts. These results imply that the overall activity level of the contrasted shoals is not a necessary condition for small shoals discrimination in angelfish. On the other hand, the results obtained for the large shoals, together with results obtained in the control treatments (equal numerical contrasts and differing activity levels), suggest that activity is a sufficient condition for discrimination when large shoals are involved. Further experiments are needed to evaluate the influence of other continuous variables, and to assess whether the mechanisms underlying performance are comparable to those suggested for other animals.
Adults, infants and non-human primates are thought to possess similar non-verbal numer-ical systems, but there is considerable debate regarding whether all vertebrates share the same numerical abilities. Despite an abundance of studies, cross-species comparison remains difficult because the methodology employed and the context of species exam-ination vary considerably across studies. To fill this gap, we used the same procedure, stimuli, and numerical contrasts to compare quantity abilities of five teleost fish: redtail splitfin, guppies, zebrafish, Siamese fighting fish, and angelfish. Subjects were trained to discriminate between two sets of geometrical figures using a food reward. Fish initially were trained on an easy numerical ratio (5 vs. 10 and 6 vs. 12). Once they reached the learning criterion, they were subjected to non-reinforced probe trials in which the set size was constant but numerical ratios varied (8 vs. 12 and 9 vs. 12).They also were subjected to probe trials in which the ratio was constant, but the total set size was increased (25 vs. 50) or decreased (2 vs. 4). Overall, fish generalized to numerosities with a 0.67 ratio, but failed with a 0.75 ratio; they generalized to a smaller set size, but not to a larger one. Only minor differences were observed among the five species. However, in one species, zebrafish, the proportion of individuals reaching the learning criterion was much smaller than in the others. In a control experiment, zebrafish showed a similar lower performance in shape discrimination, suggesting that the observed difference resulted from the zebrafish's diffi-culty in learning this procedure rather than from a cross-species variation in the numerical domain.
Zebrafish is an emerging model in the study of brain function; however, knowledge about its behaviour and cognition is incomplete. Previous studies suggest this species’ limited ability in visual learning tasks compared to other teleosts. In this study, we systematically examined zebrafish’s ability to learn to discriminate colour, shape, size, and orientation of figures using an appetitive conditioning paradigm. Contrary to earlier reports, the zebrafish successfully completed all tasks. Not all discriminations were learned with the same speed and accuracy. Subjects discriminated the size of objects better than their shape or colour. In all three tasks, they were faster and more accurate when required to discriminate between outlined figures than between filled figures. With stimuli consisting of outlines, the learning performance of zebrafish was comparable to that observed in higher vertebrates. Zebrafish easily learned a horizontal–vertical discrimination task, but like many other vertebrates, they had great difficulty discriminating a figure from its mirror image. Performance was more accurate for subjects reinforced on one stimulus (green over red, triangle over circle, large over small). Unexpectedly, these stimulus biases occurred only when zebrafish were tested with filled figures, suggesting some causal relationship between stimulus preference, learning bias and performance.
Visual stimuli can evoke complex behavioral responses, but the underlying streams of neural activity in mammalian brains are difficult to follow because of their size. Here, I review the visual system of zebrafish larvae, highlighting where recent experimental evidence has localized the functional steps of visuomotor transformations to specific brain areas. The retina of a larva encodes behaviorally relevant visual information in neural activity distributed across feature-selective ganglion cells such that signals representing distinct stimulus properties arrive in different areas or layers of the brain. Motor centers in the hindbrain encode motor variables that are precisely tuned to behavioral needs within a given stimulus setting. Owing to rapid technological progress, larval zebrafish provide unique opportunities for obtaining a comprehensive understanding of the intermediate processing steps occurring between visual and motor centers, revealing how visuomotor transformations are implemented in a vertebrate brain.
[in: Annual Review of Vision Science, Vol. 5]
The ability to use quantitative information is thought to be adaptive in a wide range of ecological contexts. For nearly a century, the numerical abilities of mammals and birds have been extensively studied using a variety of approaches. However, in the last two decades, there has been increasing interest in investigating the numerical abilities of teleosts (i.e. a large group of ray-finned fish), mainly due to the practical advantages of using fish species as models in laboratory research. Here, we review the current state of the art in this field. In the first part, we highlight some potential ecological functions of numerical abilities in fish and summarize the existing literature that demonstrates numerical abilities in different fish species. In many cases, surprising similarities have been reported among the numerical performance of mammals, birds and fish, raising the question as to whether vertebrates' numerical systems have been inherited from a common ancestor. In the second part, we will focus on what we still need to investigate, specifically the research fields in which the use of fish would be particularly beneficial, such as the genetic bases of numerical abilities, the development of these abilities and the evolutionary foundation of vertebrate number sense.
This article is part of a discussion meeting issue ‘The origins of numerical abilities’.
Two theories compete to explain how we estimate the numerosity of visual object sets. The first suggests that the apparent numerosity is derived from an analysis of more low-level features like size and density of the set. The second theory suggests that numbers are sensed directly. Consistent with the latter claim is the existence of neurons in parietal cortex which are specialized for processing the numerosity of elements in the visual scene. However, recent evidence suggests that only low numbers can be sensed directly whereas the perception of high numbers is supported by the analysis of low-level features. Processing of low and high numbers, being located at different levels of the neural hierarchy should involve different receptive field sizes. Here, I tested this idea with visual adaptation. I measured the spatial spread of number adaptation for low and high numerosities. A focused adaptation spread of high numerosities suggested the involvement of early neural levels where receptive fields are comparably small and the broad spread for low numerosities was consistent with processing of number neurons which have larger receptive fields. These results provide evidence for the claim that different mechanism exist generating the perception of visual numerosity. Whereas low numbers are sensed directly as a primary visual attribute, the estimation of high numbers however likely depends on the area size over which the objects are spread.
Noribogaine is the main psychoactive metabolite of the hallucinogenic drug ibogaine, and is a particularly interesting compound potentially useful to treat dependence and various psychiatric disorders. Here, we report the effects of noribogaine on anxiety and locomotion in zebrafish (Danio rerio), a new promising model organism in neurobehavioral and psychopharmacological research. Adult zebrafish were subjected to the 5 min novel tank test (NTT) following an acute, 20 min drug immersion in 1, 5 and 10 mg/L noribogaine. Overall, noribogaine produced robust anxiolytic-like behavior in zebrafish (increasing the time spent and transition to the top half compartment and reducing freezing bouts) without overt effects on fish locomotion. Taken together, these results indicate that noribogaine modulates the components of the acute stress response related to emotionality and anxiety behaviors, implicating this drug as a potentially useful non-sedative anxiolytic agent.
Continuous Issues in Numerical Cognition: How Many or How Much re-examines the widely accepted view that there exists a core numerical system within human beings and an innate ability to perceive and count discrete quantities. This core knowledge involves the brain's intraparietal sulcus, and a deficiency in this region has traditionally been thought to be the basis for arithmetic disability. However, new research findings suggest this wide agreement needs to be examined carefully and that perception of sizes and other non-countable amounts may be the true precursors of numerical ability. This cutting-edge book examines the possibility that perception and evaluation of non-countable dimensions may be involved in the development of numerical cognition. Discussions of the above and related issues are important for the achievement of a comprehensive understanding of numerical cognition, its brain basis, development, breakdown in brain-injured individuals, and failures to master mathematical skills. Serves as an innovative reference on the emerging field of numerical cognition and the branches that converge on this diverse topic Features chapters from leading researchers in the field Includes an overview of the multiple disciplines that comprise numerical cognition and discusses the measures that can be used in analysis Introduces novel ideas that connect non-countable continuous variables to numerical cognition
Adults’ ability to process numerical information can be traced back to the first days of life. The cognitive mechanisms underlying numerical representations are functional in preverbal infants, who are able to both track a small number of individuals and to estimate the numerosity of large sets across different modalities. This ability is closely linked to their ability to compute other quantitative dimensions such as spatial extent and temporal duration. In fact, the human mind establishes, early in life, spontaneous links between number, space, and time, which are privileged relative to links with other continuous dimensions (like loudness and brightness). Finally, preverbal infants do not only associate numbers to corresponding spatial extents but also to different spatial positions along a spatial axis. It is argued that these number–space mappings are at the origins of the “mental number line” representation, which is already functional in the first year of life.
When two food patches are available, individuals of many animal species feed on the larger one, a preference frequently used to study numerical abilities in mammals and birds. We employed this method to investigate, for the first time, food quantity discrimination and its underlying mechanisms in a fish, the guppy, Poecilia reticulata. Guppies facing two sets of similar-sized food items successfully discriminated numerosity up to a 0.5 ratio (one versus four and two versus four items, but not two versus three or three versus four food items). A further experiment suggested that guppies attended to cumulative surface area of food items rather than number to select the larger quantity. Moreover, in a two versus four discrimination in which the cumulative surface area occupied by food was matched by using larger items in the set with fewer items, guppies unexpectedly showed a preference for the smaller numerosity. Since this result might be explained by assuming that guppies selected the larger food item, we performed additional experiments to test this hypothesis. Guppies were observed to be very accurate in estimating item size, being able to discriminate between two food items that differed by a ratio of 0.75 in surface area. The attraction to the larger food item was so strong that guppies preferred the set containing the largest item even when the other set contained a double quantity of food. Since guppies in the wild forage in groups and compete for food, we hypothesized that in this species natural selection has favoured cognitive mechanisms allowing a rapid and efficient choice of the most profitable food item within the patch.
Discrimination of quantity (magnitude) was investigated in zebrafish (Danio rerio). Male zebrafish chose to approach the location previously occupied by the larger in number between 2 groups of female conspecifics (no longer visible at test) in sets of 1 versus 2 items, and 2 versus 3 items, but failed at 3 versus 4 items; similarly, when tested with larger numbers, zebrafish succeeded with 2 versus 4, 4 versus 6, and 4 versus 8 items, but failed with 6 versus 8 items. The results suggest that zebrafish rely on an approximate number system to discriminate memorized sets of conspecifics of different magnitudes, the degree of precision in recall being mainly dependent on the ratio between the sets to be discriminated. (PsycINFO Database Record
The ability to discriminate between sets of items differing in quantity has shown a growing interest in comparative studies as a diversity of animal species exhibit such quantitative competence. Previous studies with angelfish (Pterophyllum scalare) have demonstrated that this species is capable of spontaneously discriminating between fully visible groups (shoals) of conspecifics of different numerical size. In the present study, we investigated quantity discrimination in angelfish adopting a new procedure that we expected to make the task more difficult for the fish. During a pretest period, angelfish were allowed to fully see shoals of conspecifics of different numerical size, subsequently all fish but 1 in each stimulus shoal were hidden behind opaque barriers. Thus, during testing, experimental fish had to rely on their working memory, which implies a certain level of mental representation of the quantities or numbers discriminated. Angelfish chose the larger shoal with similar accuracy when 1 versus 2, 1 versus 3, 1 versus 4, 2 versus 3, and 2 versus 4 stimulus fish were contrasted, but failed to distinguish shoals when 3 versus 4, 4 versus 5, and 4 versus 6 fish were contrasted. Strong similarities were observed in relation with our previous procedure indicating the robustness of the quantity discrimination abilities of this species. Our results imply that angelfish form internal representations and demonstrate that these fish can make comparisons between small quantities of items while relying on their working memory alone. (PsycINFO Database Record (c) 2014 APA, all rights reserved).
Despite considerable research on numeric judgements in animals, uncertainty remains about both the underlying mechanisms and the role of training. To address these issues, we study quantity discrimination in jungle crows that have previously been shown to select the larger of two quantities following training. In the current study, we examined whether jungle crows are able to discriminate between different quantities of food items without prior training. Using a simultaneous two-alternative test, we studied whether their performance reveals the underlying mechanism being used for discrimination of small and large quantities in other animals. First, jungle crows were tested with a choice between two discrete homogeneous quantities; one smaller and one larger, with ratios of 0.5 (1 versus 2, 2 versus 4 and 4 versus 8), 0.67 (2 versus 3, 4 versus 6 and 8 versus 12) and 0.75 (3 versus 4, 6 versus 8 and 12 versus 16). Then, we conducted a test using a non-numerical cue where the volume of comparison stimuli was equalized. Jungle crows selected the larger of the two quantities in all comparisons, except those when both quantities were large (6 versus 8, 8 versus 12 and 12 versus 16). Furthermore, accuracy of selection of the larger quantity declined with increasing numerical magnitude. These results suggest that in a spontaneous discrimination task, jungle crows use an object-file mechanism to compare quantities, even when the number of items in one of the arrays exceeds four. The crows showed no preference for the smaller or the larger quantity when the volume cue was removed. This lack of bias may suggest an ecological role of selection for a patch with higher amounts of food and not necessarily the exact number of food items for optimal foraging in jungle crows.
The zebrafish (Danio rerio) is rapidly becoming a popular model organism in pharmacogenetics and neuropharmacology. Both larval and adult zebrafish are currently used to increase our understanding of brain function, dysfunction, and their genetic and pharmacological modulation. Here we review the developing utility of zebrafish in the analysis of complex brain disorders (including, for example, depression, autism, psychoses, drug abuse and cognitive disorders), also covering zebrafish applications towards the goal of modeling major human neuropsychiatric and drug-induced syndromes. We argue that zebrafish models of complex brain disorders and drug-induced conditions have become a rapidly emerging critical field in translational neuropharmacology research.
An investigation of the predator-evasion behaviour of minnow (Phoxinus phoxinus) shoals confronted with a pike (Esox lucius) showed that individual minnows generally chose the behaviour that minimized their chance of being eaten by the predator. As soon as the pike had been detected, minnows switched from dispersed small shoals to a single compact school. They then commenced inspection behaviour, during which individuals or groups approached the predator. This inspection served to confirm recognition of the pike and provide information on its behaviour. Avoidance and skittering behaviour took place when the pike began stalking. It was only when the predator escalated its attack and struck at the shoal that the minnows performed their most costly predator evasion tactics, such as flash expansion and fountain. After such tactics individuals often became separated from the shoal and as such were most vulnerable to capture. As a last resort, individual minnows hid among stones. Minnows from provenances with and without pike exhibited a similar repertoire of antipredator behaviour patterns, but those sympatric with the predator integrated their tactics more effectively and regained pre-exposure behaviour sooner after each encounter. Shoal size had an important effect on the execution of tactics. Minnows in shoals of 10 were more likely than minnows in shoals of 20 or 50 to abandon schooling behaviour and seek cover as individuals.
Shoaling fish are expected, in many cases, to gain fitness benefits from being in a larger shoal and previous experiments have shown that fish are indeed capable of choosing between shoals of different sizes. We investigated the influence of shoal activity on shoal size preference in the zebrafish. We gave test fish the choice between shoals of one to four stimulus fish, presented at two different water temperatures, and so differing in their activity levels. Where all stimulus fish were in water of the same temperature, test fish generally preferred the larger shoal. However, this preference could be reduced by presenting the larger shoal in colder water and so reducing its activity. We discuss these findings with reference to the factors that may influence shoal activity, the effect of temperature on shoaling behaviour and the mechanisms that may be used by fish to discriminate shoal size.
Considerable research has investigated infants' numerical capacities. Studies in this domain have used procedures of habituation, head turn, violation of expectation, reaching, and crawling to ask what quantities infants discriminate and represent visually, auditorily as well as intermodally. The concensus view from these studies is that infants possess a numerical system that is amodal and applicable to the quantification of any kind of entity and that this system is fundamentally separate from other systems that represent continuous magnitude. Although there is much evidence consistent with this view, there are also inconsistencies in the data. This paper provides a broad review of what we know, including the evidence suggesting systematic early knowledge as well as the peculiarities and gaps in the empirical findings with respect to the concensus view. We argue, from these inconsistencies, that the concensus view cannot be entirely correct. In light of the evidence, we propose a new hypothesis, the Signal Clarity hypothesis, that posits a developmental role for dimensions of continuous quantity within the discrete quantity system and calls for a broader research agenda that considers the covariation of discrete and continuous quantities not simply as a problem for experimental control but as information that developing infants may use to build more precise and robust representations of number.
Previous studies investigating quantity discrimination have shown that angelfish are able to select the larger of two groups of conspecifics (shoals). The discrimination limits shown by angelfish were similar to those found for other vertebrates when large (≥4) and small quantities (<4) were presented. However, in these studies, no attempt was made to control for non-numerical features of the stimulus shoals and thus the question whether numerical or some quantitative attributes of the shoals were utilized for making the choices could not be answered. Here, we investigate whether angelfish can discriminate between shoals differing in numerical size using non-numerical attributes. We systematically manipulate density, inter-fish distance, and overall space occupied by the shoals, one factor at a time, and analyse the choices angelfish made between the contrasting stimulus shoals. The stimulus shoals consisted of contrasts between large (10 vs. 5) and small (3 vs. 2) number of conspecifics. We found density to be a sufficient condition for discrimination between large shoals as the test subjects preferred the more dense shoal. Manipulation of inter-fish distance indicated that this variable is not a necessary factor in discrimination at either shoal size contrast. Likewise, we found that the size of space occupied by the contrasted shoals also did not significantly influence discrimination. Sensitivity to the density of large shoals indicates that angelfish can discriminate shoal size using this non-numerical cue. Nevertheless, the factors we examined may represent only a subset of all possible non-numerical features upon which angelfish may base their discrimination. Thus, we suggest that further research is required to clarify whether and under what circumstances angelfish may use numerical or non-numerical features when discriminating between shoals of differing size.
Describes a theory of temporal control which treats responding of animal Ss at asymptote under a variety of learning procedures. Ss are viewed as making estimates of the time to reinforcement delivery using a scalar-timing process, which rescales estimates for different values of the interval being timed. Scalar-timing implies a constant coefficient of variation. Expectancies of reward based on these estimates are formed, and a discrimination between response alternatives is made by taking a ratio of their expectancies. In periodic schedules of reinforcement the discrimination is between local and overall expectancy of reward. In psychophysical studies of duration discrimination, the expectancy ratio reduces the likelihood ratio, and in conjunction with the scalar property, results in a general form of Weber's law. The psychometric choice function describing preference for different amounts and delays of reinforcement also results in a form of Weber's law. (102 ref) (PsycINFO Database Record (c) 2012 APA, all rights reserved)
In two experiments, a manual search task explored 12- to 14-month-old infants’ representations of small sets of objects. In this paradigm, patterns of searching revealed the number of objects infants represented as hidden in an opaque box. In Experiment 1, we obtained the set-size signature of object-file representations: infants succeeded at representing precisely 1, precisely 2, and precisely 3 objects in the box, but failed at representing 4 (or even that 4 is greater than 2). In Experiment 2, we showed that infants’ expectations about the contents of the box were based on number of individual objects, and not on a continuous property such as total object volume. These findings support the hypothesis that infants maintained representations of individuals, that object-files were the underlying means of representing these individuals, and that object-file models can be compared via one-to-one correspondence to establish numerical equivalence.
Larger animal groups often provide greater protection from predators. An individual might therefore be expected to join the larger of two groups. To test this, we hypothesized that fathead minnows would choose to associate with the larger of two shoals and that the presence of a predatory largemouth bass would influence their shoal size choice. Individual minnows were presented with a series of choices between two shoal sizes, ranging from 1 to 28 fish, both with and without a predator present. Although responses were highly variable, minnows displayed an ability to choose between shoal sizes even when size differences were small, preferring the larger shoal whenever a size preference was shown. In the presence of a predator, minnows made quicker shoaling decisions and showed a strong tendency to avoid very small shoals.