Article

If I fits I sits: A citizen science investigation into illusory contour susceptibility in domestic cats (Felis silvestris catus)

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Abstract

A well-known phenomenon to cat owners is the tendency of their cats to sit in enclosed spaces such as boxes, laundry baskets, and even shape outlines taped on the floor. This investigative study asks whether domestic cats (Felis silvestris catus) are also susceptible to sitting in enclosures that are illusory in nature, utilizing cats’ attraction to box-like spaces to assess their perception of the Kanizsa square visual illusion. Carried out during the COVID-19 pandemic, this study randomly assigned citizen science participants Booklets of six randomized, counterbalanced daily stimuli to print out, prepare, and place on the floor in pairs. Owners observed and videorecorded their cats’ behavior with the stimuli and reported findings from home over the course of the six daily trials. This study ultimately reached over 500 pet cats and cat owners, and of those, 30 completed all of the study’s trials. Of these, nine cat subjects selected at least one stimulus by sitting within the contours (illusory or otherwise) with all limbs for at least three seconds. This study revealed that cats selected the Kanizsa illusion just as often as the square and more often than the control, indicating that domestic cats may treat the subjective Kanizsa contours as they do real contours. Given the drawbacks of citizen science projects such as participant attrition, future research would benefit from replicating this study in controlled settings. To the best of our knowledge, this investigation is the first of its kind in three regards: a citizen science study of cat cognition; a formal examination into cats’ attraction to 2D rather than 3D enclosures; and study into cats’ susceptibility to illusory contours in an ecologically relevant paradigm. This study demonstrates the potential of more ecologically valid study of pet cats, and more broadly provides an interesting new perspective into cat visual perception research.

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... own pet, has some significant potential advantages over studies conducted in a laboratory environment, such as providing larger samples and more naturalistic data by collecting the data in the animal's daily environment (Smith et al. 2021(Smith et al. , 2022Stewart et al. 2015). In addition, the ubiquity of dogs around the world also provides a wealth of opportunities for humans to observe and interact with dogs and to explore learning in non-human animals (ManyDogs Project et al. 2023). ...
... Further, with these approaches, subsequent re-coding or verification of the Animal Cognition owner's experimental procedure and inputted results is impossible because the only data experimenters have access to is input by guardians. Because of this limitation, some citizen science projects have used the second major variant, asking guardians to submit video data of the experiment to be coded by a researcher (Hecht and Spicer Rice 2015;Smith et al. 2021Smith et al. , 2022. This has the advantage of providing strict quality control on the data used. ...
... This may impact performance and overall data quality (Smith et al. 2022). There are also limited controls when guardians act as active experimenters (e.g., are actively presenting stimuli as seen in Stewart et al. (2015) vs. recording responses to stable stimuli like in Smith et al. (2021Smith et al. ( ,2022. Guardians typically receive written and/ or video instructions, which can be misunderstood. ...
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Citizen science approaches have grown in popularity over the years, partly due to their ability to reach a wider audience and produce more generalizable samples. In dogs, these studies, though, have been limited in their controls over materials or experimental protocols, with guardians typically reporting results without researcher supervision. Over two studies, we explored and validated a synchronous citizen science approach. We had dog guardians act as experimenters while being supervised by a researcher over Zoom. In study 1, we demonstrated that synchronous citizen science produced equivalent levels of performance to in-lab designs in a choice task. Consistent with past in-lab research, dogs selected a treat (vs. an empty plate) in a two-alternative forced-choice task. In study 2, we showed that Zoom methods are also appropriate for studies utilizing looking time measures. We explored dogs’ looking behaviors when a bag of treats was placed in an unreachable location, and dogs’ guardians were either attentive or inattentive while dogs attempted to retrieve the treats. Consistent with past work, dogs in the attentive condition looked at their guardian for longer periods and had a shorter latency to first look than dogs in the inattentive condition. Overall, we have demonstrated that synchronous citizen science studies with dogs are feasible and produce valid results consistent with those found in a typical lab setting.
... In applied ethology, the citizen science approach enables for investigation of domestic animals' behaviors and leverages the experience of caretakers (Smith et al., 2021). Most studies use videos, photos, surveys, notes, and even a diary of behaviors (Gillies and Clout, 2003;Fugazza et al., 2021;Laverack et al., 2021). ...
... It is known that they tend not to behave naturally and most of them will be very frightened (Amat et al., 2015). One good example is a cat cognition study carried out using citizen science approach during the Covid-19 pandemic (Smith et al., 2021). This study explored the tendency cats have to sit in enclosed spaces, such as boxes and baskets, and investigated whether cats sat in illusory enclosures, such as two-dimensional geometric shapes on the floor made with a piece of paper. ...
... In a previous study of Smith et al. (2021), which did not include a synchronous researcher and relied on participants recording themselves instead, 21 out of 30 participants did not follow instructions properly. In comparison, all 48 of our participants met the criteria for inclusion and analysis, thus demonstrating that our method ensures lower loss of individuals due to researcher's monitoring throughout the session. ...
Article
Owing to restrictions imposed by the COVID-19 pandemic, numerous animal behavior studies incorporated elements of citizen science and/or were conducted remotely through online video calls. For the current cognition study about communication of domestic cats, the authors added an element of synchronicity to the citizen science approach. Considering cats are territorial and do not usually leave their homes with their owners, Synchronous citizen science, as the authors designated their innovative approach, proved to be extremely beneficial. To adjust the method, five pilot trials (five cat-owner dyads) were conducted, followed by a subsequent sample with 48 cat-owner dyads. Internet access and a free video call app were required for participation as the experiment was to be conducted during an online video call scheduled at the most convenient time for the cat-owner dyad. Before the experiment, a script detailing the procedure to be followed was sent to the participants. During the video call, two types of interactions were recorded: feeding and playing. In the first scenario, the cat would be offered their favorite food for up to five seconds. Then, the owner would restrict access by holding the feeder for 60 s, after which they would offer the food to the cat once again. In the second scenario, the owner would initiate a playing session using a new wand toy sent to them by the authors. The owner would play with the cat for 40 s, cease interaction for 60 s, and resume playing for 40 s. The interactions lasted two and four minutes, respectively, and were monitored by the researcher, who could guide the participant in the experiment, if needed. This short communication aims at presenting this new methodology, developed and refined in the pilot sample of five dyads. After the first round of experiments discussed here, the authors replicated the procedure with a full sample of 48 dyads and achieved high success and data reliability. Prioritizing humans' and cats’ welfare during the COVID-19 pandemic called for creative research methods, which eventually led to exciting paths in terms of analyzing cat behavior in their familiar environment with minimal disturbances.
... To safely test animals during the COVID-19 pandemic, we recruited participants in a community science method identical to the procedure of G. E. Smith et al. (2021). Community science is a useful method to include participants unable to come to a research facility (e.g., fearful, or aggressive dogs), but also to test pets in an environment where they are likely to behave in a naturalistic manner. ...
... ful method to include participants unable to come to a research facility (e.g., fearful, or aggressive dogs), but also to test pets in an environment where they are likely to behave in a naturalistic manner. Community science is further ideal for studying domestic cats, as many cats become severely stressed in novel environments (Amat et al., 2016;G. E. Smith et al., 2021). ...
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Within human–animal dyadic interactions, dog–human gaze has been identified as the crux of several important visual behaviors, such as looking back, gaze-following, and participation in an oxytocin feedback loop. It has been posited that this gaze behavior may have been motivated and sustained by cooperative relationships between dogs (Canis lupus familiaris) and humans (e.g., hunting, service roles), however, to investigate why gaze evolved, a comparison to a domesticated species that lacks a protracted history of cooperative companionship is needed: the domestic cat (Felis catus). In this study, we compare the gaze duration to owners of cats and dogs in a community science setting. We replicated previous gaze studies with dogs, wolves (Nagasawa et al., 2015), and dingoes (Johnston et al., 2017), requesting owners to sit with their pets for 5 min and interact as they normally would. Cats and dogs gazed at their owners for similar durations, but durations of petting and physical contact were significantly lower with cats. Gaze correlated significantly with vocalizations in dogs; however, no other correlations were significant. Dogs gazed less in our community science setting than dogs tested previously in-lab (Nagasawa et al., 2015). Ultimately, cats resemble dogs in their general gaze patterns, but not in most interactions with their owner. Future research should aim to include feral cats or wild cat species to shed light on gaze behavior development in the genus, while more community science work can identify the behaviors that shift for dogs between familiar and unfamiliar environments.
... An alternative experimental method involves spontaneous choice, in which an individual exhibits attraction to an illusion due to its biological relevance (most often food). A study by Smith et al. (2020) [10] re-examined domestic cats' perception of the Kanizsa square by presenting cats with the illusion, a square, and a control, relying on cats' attraction to enclosed spaces to determine the illusory perception. The authors found that the subjects stood/sat in the illusion as much as in the square and more than in the control, confirming that cats demonstrate a human-like susceptibility to the illusion. ...
... This four-month study took place from May to September 2021 and was adapted from Regaiolli et al.'s (2019) [17] methodology into a community science paradigm due to its efficacy in examining non-human animal behavior and cognition (e.g., [10,22,23] The three stimuli presented were: Rotating Snakes; a control stimulus with the same pattern as the illusion that does not elicit perceived motion (hereafter called Plain Snakes); and a control stimulus consisting of a simple pattern of overlapping circles that also does not elicit perceived motion (hereafter called Big Circles) ( Figure 1). The Rotating Snakes illusion consists of alternating black, blue, white, and yellow blobs (black, dark grey, white, light grey for the achromatic version). ...
Article
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Illusions—visual fields that distort perception—can inform the understanding of visual perception and its evolution. An example of one such illusion, the Rotating Snakes illusion, causes the perception of motion in a series of static concentric circles. The current study investigated pet dogs’ and cats’ perception of the Rotating Snakes illusion in a community science paradigm. The results reveal that neither species spent significantly more time at the illusion than at either of the controls, failing to indicate susceptibility to the illusion. Specific behavioral data at each stimulus reveal that the most common behaviors of both species were Inactive and Stationary, while Locomotion and Pawing were the least common, supporting the finding that susceptibility may not be present. This study is the first to examine susceptibility to the Rotating Snakes illusion in dogs, as well as to directly compare the phenomenon between dogs and cats. We suggest future studies might consider exploring alternative methods in testing susceptibility to motion illusions in non-human animals.
... The cat (Felis catus) is a species that retained its original hunting skills as a domesticated mesopredator, 8 however, it also widely shares the anthropogenic niche with humans as one of the most successful companion animals. 9 There is accumulating evidence about cats' capacity for complex cognition, for example, they performed successfully in the so-called ''Kanizsa-square'' paradigm 10 ; and cats were also capable of spontaneous quantity-discrimination (adult cats, 11 kittens 12 ). Moreover, perhaps as a sign of evolutionary adaptation to their new social environment, cats show remarkable performance in many tasks involving social cognition: they can follow human visual signals, such as gazing, 13 respond to ostensive cues, 14 and prefer cat-directed speech patterns. ...
Article
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Various animal species can make a priori decisions about the passability of openings, based on their own size knowledge. So far no one has tested the ability for self-representation in cats. We hypothesized that cats may rely on their size awareness when they have to negotiate small openings. Companion cats (N = 30) were tested with incrementally decreasing sized openings, which were either the same height, or the same width. Cats approached and entered even the narrowest openings, but they slowed down before reaching, and while passing through the shortest ones. Because of their specific anatomical features and cautious locomotory strategy, cats readily opt for the trial-and-error method to negotiate narrow apertures, but they seemingly rely on their body-size representing capacity in the case of uncomfortably short openings. Ecologically valid methodologies can provide answers in the future as to whether cats would rely on their body awareness in other challenging spatial tasks.
... For example, Kanizsa triangles (Figure 3b) produce the illusion of a bright triangle via the use of fragmented Pac-man-like stimuli, none of which fully outline the perceived shape. Research in nonhuman animals has revealed that illusory contours such as Kanizsa figures are perceived by cats (Smith et al. 2021), mice (Okuyama-Uchimura & Komai 2016), goldfish (Wyzisk & Neumeyer 2007), cuttlefish (Zylinski et al. 2012), and bees (Horridge et al. 2022) but not rats (Minini & Jeffery 2006). This taxonomic diversity suggests a general benefit of heuristics that support shape detection and boundary completion during object perception, perhaps especially in spatially complex environments where shape-diagnostic edges may be interrupted by visual obstructions (Peterhans & Von Der Heydt 1991). ...
Article
Animals live in visually complex environments. As a result, visual systems have evolved mechanisms that simplify visual processing and allow animals to focus on the information that is most relevant to adaptive decision making. This review explores two key mechanisms that animals use to efficiently process visual information: categorization and specialization. Categorization occurs when an animal's perceptual system sorts continuously varying stimuli into a set of discrete categories. Specialization occurs when particular classes of stimuli are processed using distinct cognitive operations that are not used for other classes of stimuli. We also describe a nonadaptive consequence of simplifying heuristics: visual illusions, where visual perception consistently misleads the viewer about the state of the external world or objects within it. We take an explicitly comparative approach by exploring similarities and differences in visual cognition across human and nonhuman taxa. Considering areas of convergence and divergence across taxa provides insight into the evolution and function of visual systems and associated perceptual strategies.
... The use of a video recorded by a small video camera attached to the collar around the neck of the pet. video-based approach for analysing cat-cat interactions and dynamics, although the methodology has been used for other research topics in cat behavioural research including cognition (Smith et al., 2021) and communication (Fukimoto et al., 2023). Videos recorded by lay people have also been used in behavioural research in other species, including dogs (e.g. ...
Article
Citizen science, which involves engaging the general public in research tasks, is increasingly used in animal behaviour studies. In this review we conducted a SWOT analysis (Strengths, Weaknesses, Opportunities, Threats) to evaluate two methodologies of data collection using citizen science in order to study cat-cat interactions: online survey responses and caregiver-recorded home videos analysed by researcher(s). Using the SWOT-analysis on both methodologies, we listed intrinsic aspects that facilitate (Strengths) or interfere with (Weaknesses) reaching scientific goals, as well as the features that the methodology may be able to capitalise on (Opportunities) or which limit its value (Threats). A major strength of online surveys is the possibility to access caregivers’ specific knowledge of their cats, while sampling bias often is a potential weakness. Opportunities of surveys are the methodology´s flexibility and data collection efficiency, but at the same time suffering from threats related to biases associated with caregiver interpretation of their pet’s behaviour. Strengths of caregiver-recorded videos capturing cats’ behaviour include that they allow expert behavioural observations and scoring in a systematic manner (e.g. using an ethogram) and thus yielding quantitative data (whose reliability can be tested between and within observers). Furthermore, given the ubiquity of smartphones, filming cats is not a burden for most caregivers, and the collected recordings can potentially contain high-quality data that may otherwise be inaccessible, or subject to bias if a researcher had been present in the home environment. Though, caregivers’ influence on and lack of standardisation of the recordings are weaknesses which possibly influence the quality of the collected data. Opportunities include public engagement with science, while possible Threats may be related to privacy of the caregivers participating. In this review we consider in more detail each of the four SWOT components related to each methodology in order to optimise cat behaviour research in the future. The authors suggest strategies for future studies using the research methodologies discussed in this review and give specific recommendations when using caregiver-recorded videos in behavioural studies. Additionally, smart combinations of both online surveys with home videos recorded by caregivers might overcome some limitations of the individual methodologies, and would thus be a potentially stronger approach.
... This relates mostly to circles 1 and 2, and somewhat to circle 3 in fig. 3 (Bird et al., 2014;Johnston et al., 2022), in general, citizen science data has been invaluable for ecological research (e.g. Border et al., 2017;Gordo et al., 2021;Greenwood, 2007;Smith et al., 2021;Sweet, Rödl, et al., 2022). The first study of this thesis is one such investigation that uses public citizen science data in addition to government-managed data. ...
Thesis
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This thesis aims to assess and add to the basic ecological and social-ecological knowledge within urban ecosystems. Human attitudes strongly influence the ecology of cities, and this in turn influences how people that live in cities perceive nature in cities. I addressed basic questions relating to animal biodiversity in German cities and attitudes towards the animals that are present in them. These questions relate to the apparent paucity of information on these topics in Germany, and research towards urban nature as a whole. Link to thesis: https://mediatum.ub.tum.de/?id=1695254
... This was probably because food rewards seemed insufficient to motivate the laboratory dogs to participate in the task, and/or their willingness to participate could have been influenced by a variety of situational factors, such as the unfamiliarity of the situation and/or the proximity of the stranger. The finding that one of the principal components, Willingness to interact, was composed of the dogs' willingness to accept food as well as behavioral variables coded in the abovementioned three tasks, clearly indicates that the degree of individual willingness to participate is one of the key challenges in testing laboratory dogs [similar to some other animals, like cats; e.g., Smith et al. (33)]. ...
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For a long time, oxytocin has been thought to have a generally positive effect on social cognition and prosocial behavior; however, recent results suggested that oxytocin has beneficial effects only under certain conditions. The aim of the present study was to explore potential associations between social competence and the effect of intranasal oxytocin on the social behavior of laboratory beagle dogs. We expected oxytocin treatment to have a more pronounced positive effect on dogs with lower baseline performance in a social test battery. Thirty-six adult dogs of both sexes received 32 IU intranasal oxytocin and physiological saline (placebo) treatment in a double-blind, cross-over design, with 17–20 days between the two sessions. Forty minutes after the treatment, dogs participated in a social test battery consisting of eight situations. The situations were carried out within one session and took 20–30 min to complete. Principal component analysis on the coded behaviors identified four components (Willingness to interact, Preference for social contact, Non-aversive response to nonsocial threat, and Non-aversive response to social threat). The subjects' behavior during the placebo condition was used to assess their baseline performance. We found that oxytocin treatment had a differential effect on the behavior depending on the baseline performance of the individuals in all components, but only two treatment × baseline performance interactions remained significant in a less sensitive analysis. In accordance with our hypothesis, oxytocin administration increased dogs' contact seeking and affiliative behaviors toward humans but only for those with low baseline performance. Dogs with low baseline performance also showed significantly more positive (friendly) reactions to social threat after oxytocin administration than after placebo, while for dogs with high baseline performance, oxytocin administration led to a more negative (fearful) reaction. These results indicate that similar to those on humans, the effects of oxytocin on dogs' social behavior are not universally positive but are constrained by individual characteristics and the context. Nevertheless, oxytocin administration has the potential to improve the social behavior of laboratory beagle dogs that are socially less proficient when interacting with humans, which could have both applied and animal welfare implications.
... We see the advent of the era when high quality video and sound recording became possible with the extremely widespread handheld devices for basically everyone, and large files can be easily transmitted through the internet. Consequently, amateurs can be involved in increasingly sophisticated and interactive research protocols, where they can be instructed by the researchers how and what to do while conducting experiments with, for example, their companion animals at home (Laverack et al., 2021;Smith et al., 2021). ...
... Among these, Tiira (2021) used a citizen science approach to test a novel device to combat separation anxiety in dogs. Smith et al. (2021) brought us their widely applauded study on cats' performance in the so-called illusory contour susceptibility paradigm. This paper was downloaded over 26,000 times since its publication in July 2021. ...
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The comparative study of the perception of visual illusions between different species is increasingly recognized as a useful noninvasive tool to better understand visual perception and its underlying mechanisms and evolution. The aim of the present study was to test whether the domestic cat is susceptible to the Delboeuf illusion in a manner similar to other mammalian species studied to date. For comparative reasons, we followed the methods used to test other mammals in which the animals were tested in a 2-way choice task between same-size food stimuli presented on different-size plates. In 2 different control conditions, overall the 18 cats tested spontaneously chose more often the larger amount of food, although at the individual level, they showed interindividual differences. In the Delboeuf illusion condition, where 2 equal amounts of food were presented on different-size plates, all cats chose the food presented on the smaller plate more often than on the larger one, suggesting that they were susceptible to the illusion at the group level, although at the individual level none of them performed significantly above chance. As we found no correlation between the cats’ overall performance in the control conditions and their performance in the illusion condition, we propose that the mechanisms underlying spontaneous size discrimination and illusion perception might be different. In the discussion, we compare the results of the present study with the results for other previously tested mammals and highlight some possible reasons for their similarities and differences.
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Family dogs and dog owners offer a potentially powerful way to conduct citizen science to answer questions about animal behavior that are difficult to answer with more conventional approaches. Here we evaluate the quality of the first data on dog cognition collected by citizen scientists using the Dognition.com website. We conducted analyses to understand if data generated by over 500 citizen scientists replicates internally and in comparison to previously published findings. Half of participants participated for free while the other half paid for access. The website provided each participant a temperament questionnaire and instructions on how to conduct a series of ten cognitive tests. Participation required internet access, a dog and some common household items. Participants could record their responses on any PC, tablet or smartphone from anywhere in the world and data were retained on servers. Results from citizen scientists and their dogs replicated a number of previously described phenomena from conventional lab-based research. There was little evidence that citizen scientists manipulated their results. To illustrate the potential uses of relatively large samples of citizen science data, we then used factor analysis to examine individual differences across the cognitive tasks. The data were best explained by multiple factors in support of the hypothesis that nonhumans, including dogs, can evolve multiple cognitive domains that vary independently. This analysis suggests that in the future, citizen scientists will generate useful datasets that test hypotheses and answer questions as a complement to conventional laboratory techniques used to study dog psychology.
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We present the first evidence that cats experience visual illusions and that a non-human animal can see illusory motion. In three videos we show cats reacting with hunting behavior when watching the Rotating Snakes illusion. This is taken to mean that cats see illusory motion in this image due to the propensity of cats to pursue movement. This is further supported by a survey where 29% of the respondents answered that their cat reacted to the illusion. A number of preferential looking experiments were also indicative of cats experiencing the illusion, but not conclusively so.
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The human visual system sees an illusory contour where there is a fault line across a regular striped pattern. We demonstrate that bees respond as if they see the same illusory contour. There is also a type of neuron in the lobula of the dragonfly optic lobe which responds directionally to motion of the illusory contour as if to an edge or line. Apparently insects have a mechanism that sees illusory contours and therefore assists in the demarcation of edges and objects at places where local contrast falls to zero at an edge, or where one textured object partially obscures another. These results suggest that insect vision, although spatially crude and low in processing power, sees separate objects by similar mechanisms to our own.
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This paper reports a cross-cultural investigation of susceptibility to geometrical illusions based upon student samples in Ghana and Scotland. The results indicate that: (1)cultural differences in susceptibility to illusions can be found among Ss exposed to lengthy formal education of a similar type; (2) the perceptual consequences of professional training in art and architecture tend to be moderate within a culture; but (3) the combined outcome of training across cultures tends to reduce cultural differences in susceptibility to illusions. They also raise a problem concerning the supposed positive relationship between field dependence and susceptibility to illusions. Some implications of these results are considered.
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The processing of Kanizsa-square illusory figures was studied in two experiments with four humans and two chimpanzees. Subjects of the two species were initially trained to select a Kanizsa-square illusory figure presented in a computerized two-alternative forced choice task. After training, adding narrow closing segments to the pacman inducers that composed the Kanisza illusory figures lowered performance in both chimpanzees and humans, suggesting that the discrimination could be controlled by the perception of illusory forms. A second experiment assessed transfer of performance with five sets of figures in which the size of the inducers and their separation were manipulated. Only for chimpanzees was performance directly controlled by separation, suggesting that chimpanzees are more sensitive than humans to the separation between visual elements.
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The present study examined whether color spreading and illusory contours in the neon color spreading effect of Ehrenstein figures are governed by different mechanisms. In the experiment, Ehrenstein figures with colored crosses inserted in the central gaps were used. There were three luminance conditions: the luminance of the Ehrenstein figures was lower than, the same as, or higher than the luminance of the background. In each condition, 16 trials (2 sets of instructions X 8 repetitions) were conducted in a random order. Subjects were required to adjust the luminance of the colored crosses according to one of the two sets of instruction given before each trial. One was to adjust the upper and lower thresholds in the luminance of the colored crosses such that their color was seen to spread out of the crosses. The other was to adjust the thresholds such that circular illusory contours were visible. It was found that illusory contours disappeared and the color spreading remained when the crosses and the Ehrenstein figures were in or nearly in isoluminance or when the Ehrenstein figures and the background were in isoluminance. These results suggest that color spreading and illusory contours are governed by different mechanisms.
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Robust form perception and underlying neuronal mechanisms require generalized representation of object boundaries, independent of how they are defined. One visual ability essential for form perception is reconstruction of contours absent from the retinal image. Here we show that barn owls perceive subjective contours defined by grating gaps and phase-shifted abutting gratings. Moreover, single-neuron recordings from visual forebrain (visual Wulst) of awake, behaving birds revealed a high proportion of neurons signaling such subjective contours, independent of local stimulus attributes. These data suggest that the visual Wulst is important in contour-based form perception and exhibits a functional complexity analogous to mammalian extrastriate cortex.
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This review article illustrates that mammals, birds and insects are able to perceive illusory contours. Illusory contours lack a physical counterpart, but monkeys, cats, owls and bees perceive them as if they were real borders. In all of these species, a neural correlate for such perceptual completion phenomena has been described. The robustness of neuronal responses and the abundance of cells argue that such neurons might indeed represent a neural correlate for illusory contour perception. The internal state of an animal subject (i.e., alert and behaving) seems to be an important factor when correlating neural activity with perceptual phenomena. The fact that the neural network necessary for illusory contour perception has been found in relatively early visual brain areas in all tested animals suggests that bottom-up processing is largely sufficient to explain such perceptual abilities. However, recent findings in monkeys indicate that feedback loops within the visual system may provide additional modulation. The detection of illusory contours by independently evolved visual systems argues that processing of edges in the absence of contrast gradients reflects fundamental visual constraints and not just an artifact of visual processing.
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The Ebbinghaus (Titchener) illusion was examined in a remote culture (Himba) with no words for geometric shapes. The illusion was experienced less strongly by Himba compared with English participants, leading to more accurate size contrast judgments in the Himba. The study included two conditions of inducing stimuli. The illusion was weaker when the inducing stimuli were dissimilar (diamonds) to the target (circle) compared with when they were similar (circles). However, the illusion was weakened to the same extent in both cultures. It is argued that the more accurate size judgments of the Himba derive from their tendency to prioritize the analysis of local details in visual processing of multiple objects, and not from their impoverished naming.
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Domestic dogs play many vital roles in human lives; however, relatively little is known about how they perceive the world visually. Given dogs’ recent popularity as a subject in cognitive and behavioural studies, it is important to understand how they visually interpret the world around them. One way to evaluate perception is to assess illusion susceptibility; specifically, how visual information is processed, interpreted and modified post‐retinally. While illusion susceptibility has been used across a variety of species to comparatively assess the similarities and differences in visual processing and perception, this relatively novel methodological approach has only recently been adapted to evaluate perception in domestic dogs. Here, we present a comprehensive overview of the findings from studies that have evaluated domestic dogs’ illusion susceptibility, highlighting the relevance of these results for those studying illusion susceptibility in animals as well as canine behaviour and cognition. More specifically, the ultimate goal of this review is to answer the questions: (a) Are dogs susceptible to visual illusions? (b) If so, are they susceptible to illusions in a way that parallels humans and/or other animals? (c) Are findings, within dogs, consistent and if not, how might these be interpreted and explained?
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The domestic cat (Felis silvestris catus) has shared an intertwined existence with humans for thousands of years, living on our city streets and in our homes. Yet, little scientific research has focused on the cognition of the domestic cat, especially in comparison with human's other companion, the domestic dog (Canis lupus familiaris). This review surveys the current status of several areas of cat cognition research including perception, object permanence, memory, physical causality, quantity and time discrimination, cats' sensitivity to human cues, vocal recognition and communication, attachment bonds, personality, and cognitive health. Although interest in cat cognition is growing, we still have a long way to go until we have an inclusive body of research on the subject. Therefore, this review also identifies areas where future research must be conducted. In addition to the scientific value of future work in this area, future research on cat cognition could have an important influence on the management and welfare of pet and free-roaming cats, leading to improved human-cat interactions.
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Domestic cats are exposed to a variety of stressful stimuli, which may have a negative effect on the cats' welfare and trigger a number of behavioural changes. Some of the stressors most commonly encountered by cats include changes in environment, inter-cat conflict, a poor human-cat relationship and the cat's inability to perform highly motivated behaviour patterns. Stress is very likely to reduce feed intake, and stress-related anorexia may contribute to the development of potentially serious medical conditions. Stress also increases the risk of cats showing urine marking and some forms of aggression, including redirected aggression. A number of compulsive disorders such as over-grooming may also develop as a consequence of stressful environments. Some of the main strategies to prevent or reduce stress-related behavioural problems in cats are environmental enrichment, appropriate management techniques to introduce unfamiliar cats to each other and the use of the synthetic analogue of the feline facial pheromone. As the stress response in cats depends, to a large extent, on the temperament of the animal, breeding and husbandry strategies that contribute to the cat developing a well-balanced temperament are also very useful. © The Author(s) 2015.
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Sixteen domestic cats were used to investigate the pituitary-adrenal, pituitary-gonadal and behavioral consequences of an unpredictable handling and husbandry routine. After a 10-day baseline period of standard laboratory procedures, eight cats (‘stressed cats’, STR) were subjected to a 21-day period of altered caretaking characterized by irregular feeding and cleaning times, absence of talking and petting by humans, and daily unpredictable manipulations. Eight control cats (CON group) were maintained for 21 days on the standard caretaking schedule. Behavior was recorded on time-lapse video 24 h day−1, urine was collected daily for cortisol analyses, and hormone stimulation tests with synthetic adrenocorticotropic hormone (ACTH) and luteinizing hormone-releasing hormone (LHRH) were conducted before and after the 3-week treatment period. Results indicate that the STR cats were chronically stressed by the altered caretaking routine. Urinary cortisol concentrations were consistently elevated throughout the 3-week period, adrenal sensitivity to ACTH was enhanced and pituitary sensitivity to LHRH was reduced. Active exploratory and play behavior was suppressed, and STR cats spent more time awake/alert and attempting to hide. Hiding was negatively correlated with cortisol concentration and, therefore, may be an important behavior for coping with uncontrollable and unpredictable captive environments. These results indicate that qualitatively poor caretaking is a potent psychological stressor for confined cats that may eventually compromise reproduction through behavioral or physiological mechanisms. To promote well-being, caged cats should be provided with appropriate places for concealment.
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Presents an introductory text on the 3 major areas of visual perception: sensory organization, information processing, and spatial organization. Major topics include theoretical and empirical evidence concerning the mechanisms for perceiving color, form, and movement. (PsycINFO Database Record (c) 2012 APA, all rights reserved)
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Notes that normally the visual experience of a line or a margin corresponds to a jump or a sharp discontinuity in the intensity, wave length, or spectral composition of the radiation that stimulates 2 contiguous areas of the retina. It is not, however, difficult to create situations in which there is not such correspondence. That is, quite visible contours can appear in parts of the visual field where the stimulation is completely homogeneous. These contours without gradients have lately been diversely interpreted. In this article these interpretations are discussed, and some new observations are presented. (22 ref) (PsycINFO Database Record (c) 2012 APA, all rights reserved)
Article
We report four experiments in which the strength ofedge-interpoiat-ion in illusory figure displays was tested. In Experiment 1, we investigated the relative contributions of the lengths of luminance-specified edges and the gaps between them to perceived boundary clarity as measured by using a magnitude estimation procedure. The contributionaoLthese variables were found to be best characterized by a ratio of the length of luminance-specified contour to the length of the entire edge (specified plus interpolated edge). Experiment 2 showed that this ratio predicts boundary clarity for a wide range of ratio values and display sizes.There was no evidence that illusory figure boundaries are clearer in displays with small gaps than they are in displays with larger gaps and equivalent ratios. In Experiment 3, using a more sensitive pairwise comparison paradigm, we again found no such effect. Implications for boundary interpolation in general, including perception of partially occluded objects, are discussed. The dependence of interpolation on the ratio of physically specified edges to total edgelength has thedesirable eeological consequence that unit formation will not change with variations in viewing distance.
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Fourteen adult female domestic cats were watched by two observers for 3 months. Ratings of 18 aspects of each cat's behavioural style were obtained independently from each observer. Correlations between observers were statistically significant for 15 of the 18 aspects and seven of the correlation coefficients were greater than 0·7. The ratings were compared with results of direct recording methods, where equivalent measures were available and, in five out of six cases, the results of the ratings and direct methods were significantly correlated. The rating method is, therefore, generally reliable and can be adequately validated. Some assessments of observer ratings which are not obviously and easily related to direct recordings may prove particularly useful in developmental studies of alternative modes of behaviour and the origins of individual differences.
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A new study has found that the tuning properties of neurons in the primary visual cortex of cats change as they learn an orientation-discrimination task, casting new light on the neuronal basis of perceptual learning.
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Perceptual learning has been documented in adult humans over a wide range of tasks. Although the often-observed specificity of learning is generally interpreted as evidence for training-induced plasticity in early cortical areas, physiological evidence for training-induced changes in early visual cortical areas is modest, despite reports of learning-induced changes of cortical activities in fMRI studies. To reveal the physiological bases of perceptual learning, we combined psychophysical measurements with extracellular single-unit recording under anesthetized preparations and examined the effects of training in grating orientation identification on both perceptual and neuronal contrast sensitivity functions of cats. We have found that training significantly improved perceptual contrast sensitivity of the cats to gratings with spatial frequencies near the "trained" spatial frequency, with stronger effects in the trained eye. Consistent with behavioral assessments, the mean contrast sensitivity of neurons recorded from V1 of the trained cats was significantly higher than that of neurons recorded from the untrained cats. Furthermore, in the trained cats, the contrast sensitivity of V1 neurons responding preferentially to stimuli presented via the trained eyes was significantly greater than that of neurons responding preferentially to stimuli presented via the "untrained" eyes. The effect was confined to the trained spatial frequencies. In both trained and untrained cats, the neuronal contrast sensitivity functions derived from the contrast sensitivity of the individual neurons were highly correlated with behaviorally determined perceptual contrast sensitivity functions. We suggest that training-induced neuronal contrast gain in area V1 underlies behaviorally determined perceptual contrast sensitivity improvements.
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The ability of fish to perceive subjective (or illusory) contours, ie contours that lack a physical counterpart in terms of luminance contrast gradients, was investigated. In the first experiment, redtail splitfins (Xenotoca eiseni), family Goodeidae, were trained to discriminate between a geometric figure (a triangle or a square) on various backgrounds and a background without any figure. Thereafter, the fish performed test trials in which illusory squares or triangles were obtained by (i) interruptions of a background of diagonal lines, (ii) phase-shifting of a background of diagonal lines, and (iii) pacmen spatially arranged to induce perception of Kanizsa subjective surfaces. In all three conditions, fish seemed to generalise their responses to stimuli perceived as subjective contours by humans. Fish chose, correctly, squares or triangles made of interrupted or phase-shifted diagonal lines from uniform backgrounds of diagonal lines, as well as illusory square or triangle Kanizsa figures from figures in which the inducing pacmen were scrambled. In the second experiment, fish were trained to discriminate between a vertical and a horizontal bar with luminance contrast gradients, and then tested with vertically and horizontally oriented illusory bars, created either through interruption or spatial phase-shift of inducing diagonal lines. Fish appeared to be able to generalise the orientation discrimination to illusory contours. These results demonstrate that redtail splitfins are capable of perceiving illusory contours.
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We review the evolution of domestic animals, emphasizing the effect of the earliest steps of domestication on its course. Using the first domesticated species, the dog (Canis familiaris), for illustration, we describe the evolutionary peculiarities during the historical domestication, such as the high level and wide range of diversity. We suggest that the process of earliest domestication via unconscious and later conscious selection of human-defined behavioral traits may accelerate phenotypic variations. The review is based on the results of a long-term experiment designed to reproduce early mammalian domestication in the silver fox (Vulpes vulpes) selected for tameability or amenability to domestication. We describe changes in behavior, morphology and physiology that appeared in the fox during its selection for tameability, which were similar to those observed in the domestic dog. Based on the data of the fox experiment and survey of relevant data, we discuss the developmental, genetic and possible molecular genetic mechanisms underlying these changes. We ascribe the causative role in evolutionary transformation of domestic animals to the selection for behavior and to the neurospecific regulatory genes it affects.
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Allowing very young kittens to see with only one eye at a time greatly reduces the proportion of binocular cortical cells. Compared to normal cats these specially reared animals suffer deficits in binocular depth perception while retaining normal activity in the two eyes. Evidently, binocular cells play a crucial role in stereopsis.
Article
We present the first evidence that a non-human species (the cat) is able to discriminate the orientation of illusory contours. Following Vogels and Orban45, we used two types of illusory contours. In one type, the illusory contour was defined by a number of contour-inducing semicircles, of which the endpoints were separated by a gap. In the other pattern, the inducing semicircles were shifted in phase along their diameter and their endpoints were aligned along the contour. Just noticeable differences in orientation were measured (at the 73.5% correct level), using a Wetherill and Levitt49 staircase procedure. Values in the order of 11 degrees were obtained when using the first type of illusory contour. Just noticeable differences with the second type were in the order of 17 degrees. Reducing the salience of the illusory contour, whether by scrambling the contour, or by decreasing the number or the contrast of inducing semicircles, systematically increased discrimination thresholds.
Article
Behavioural techniques were used to determine whether cats are able to see subjective contours. Through several stages of testing with increasingly complex displays, cats continued to respond to a figure defined by subjective contours. This result provides the first direct evidence that a nonhuman perceives subjective contours.
Article
We investigated responses of neurones in cortical areas 17 and 18 and in the dorsal lateral geniculate nucleus (dLGN) of the cat to a phase shift in a moving line pattern forming a border without a luminance gradient ("subjective contour"). In both areas 17 and 18, S cells and B cells respond only slightly or not at all along the phase shift while C cells respond strongly. The response of C cells is strongest for line patterns with medium line separation and decreases with smaller and larger separation. In the dLGN the relative magnitude of neuronal responses along a phase shift is similar to that of C cells. However, C cells respond uniformly along the entire phase shift, whereas geniculate cells merely respond to individual line ends along the phase shift. In addition we compared responses along a phase shift and those to a luminance gradient formed by a dotted line whose dots were separated by the same distance as the line ends along the phase shift. S cells and B cells respond preferentially to dotted lines whereas C cells and geniculate cells respond equally well along both phase shifts and dotted lines. Possible explanations for these results in terms of receptive field structure and differences in inhibitory input to the cells are discussed. Differential neurone responses may account for the perceptual distinctness of the contours with and without luminance gradients.
Article
Two separate traditions of research into the Müller-Lyer illusion have existed for at least 70 years: the ecological and the developmental. To assess the ecological hypothesis, a sufficient range in visual ecology typically has been sought in cross-cultural comparisons; however, many of these comparisons have been inconclusive, especially when other ethnic variables have been held constant. These insignificant findings are considered to result from a confounding of the ecological and developmental variables. Appropriate sampling from Temne and Eskimo populations (enabling, in turn, one variable to be matched while the other is varied) demonstrates this confounding within cultures and, when eliminated, the significant influence of each variable on susceptibility to the Müller-Lyer illusion. Further work, however, is needed to discover other factors which will account for variance in susceptibility between cultures.
Article
The question of whether phenomena of human perception such as amodal completion are also present in visual perception in the mouse was investigated. Three experiments based on a simultaneous-discrimination task were carried out with the aid of a jumping-stand similar to the one used by Lashley. The subjects were 36 male mice (DBA/J); 12 took part in each experiment. After reaching the learning criterion (87% correct responses), the mice performed 15 test trials which were interpolated between the learning trials. The test trials were constructed so as to allow the observation of any perceptual phenomena similar to amodal perception. Of the 36 subjects, 23 reached the criterion necessary to access the test phase. A significant number of them (17 out of 23, chi 2 = 5.261, p = 0.021) adopted a decisional strategy that seemed to indicate the occurrence of phenomena similar to amodal completion, and a trend in the same direction was observed in the performance of the other 6 subjects. The results allow the conclusion that a perceptual phenomenon which is similar to amodal completion in humans is present even at this phylogenetic level. Alternative interpretations are discussed.
Article
Data obtained by the author and his colleagues on deficits in learning in cats reared in the lab breeding colony are reviewed. As compared with cats who lived free in a farm during the first 3-4 months of life, the lab-reared cats showed a dramatic impairment of delayed response learning to visual and auditory stimuli. They also showed some impairment of visual discrimination learning: they commited more errors than farm-reared cats when a discrimination apparatus had a partition between the stimuli, their reaction times were longer, and after lesions of the superior colliculus-pretectum complex their relearning was impaired. Thus, even moderate impoverishment of environment can considerably affect later learning ability.
Article
A neural network model is developed to explain how visual thalamocortical interactions give rise to boundary percepts such as illusory contours and surface percepts such as filled-in brightnesses. Top-down feedback interactions are needed in addition to bottom-up feed-forward interactions to simulate these data. One feedback loop is modeled between lateral geniculate nucleus (LGN) and cortical area V1, and another within cortical areas V1 and V2. The first feedback loop realizes a matching process which enhances LGN cell activities that are consistent with those of active cortical cells, and suppresses LGN activities that are not. This corticogeniculate feedback, being endstopped and oriented, also enhances LGN ON cell activations at the ends of thin dark lines, thereby leading to enhanced cortical brightness percepts when the lines group into closed illusory contours. The second feedback loop generates boundary representations, including illusory contours, that coherently bind distributed cortical features together. Brightness percepts form within the surface representations through a diffusive filling-in process that is contained by resistive gating signals from the boundary representations. The model is used to simulate illusory contours and surface brightness induced by Ehrenstein disks, Kanizsa squares, Glass patterns, and café wall patterns in single contrast, reverse contrast, and mixed contrast configurations. These examples illustrate how boundary and surface mechanisms can generate percepts that are highly context-sensitive, including how illusory contours can be amodally recognized without being seen, how model simple cells in V1 respond preferentially to luminance discontinuities using inputs from both LGN ON and OFF cells, how model bipole cells in V2 with two colinear receptive fields can help to complete curved illusory contours, how short-range simple cell groupings and long-range bipole cell groupings can sometimes generate different outcomes, and how model double-opponent, filling-in and boundary segmentation mechanisms in V4 interact to generate surface brightness percepts in which filling-in of enhanced brightness and darkness can occur before the net brightness distribution is computed by double-opponent interactions.
Article
Visual perception of objects depends on segmentation and grouping processes that act on fragmentary input. This paper gives a brief overview of these processes. A simple geometry accounting for contour interpolation is described, and its applications to 2D, 3D, and spatiotemporal object interpolation processes are considered. A method is described for distinguishing interpolation based on this geometry from more global or top-down influences. Results suggest a separation between interpolation based on relatively local stimulus relations, which give rise to precise boundary representations, and processes involving recognition from partial information, which do not. Aspects of the model-especially the unified treatment of illusory and occluded objects-raise questions about the nature of seeing. Although it is often believed that illusory objects are perceived, while occluded objects are inferred, I suggest that both research and theory converge in supporting a more unified account. Illusory and occluded contours and surfaces do not divide into the real, the perceived, and the inferred, but are all represented, and in key respects, derive from identical perceptual processes.
Article
We used a preferential looking technique to investigate the effect of support ratio (a ratio of the physically specified contours to the total edge length) on the perception of Kanizsa illusory contours in infants aged 3-8 months. Previous work has shown that for adult observers the illusory-contour strength increases proportionally with the support ratio. When the support ratio was relatively high (66%), infants preferred illusory contours to non-illusory figures by 3-4 months of age (experiment 1). In contrast, only infants 7-8 months old showed this preference for illusory contours when the support ratio was reduced to 37% (experiment 3). Further, infants showed no preference for an outline version of the illusory-contour figure, which produced no illusory contours (experiment 2). This result confirms that the infants' preference reflects their perception of illusory contours. Our results show that (i) illusory-contour perception emerges at around 3-4 months of age, but (ii) that this ability is very limited until around 7-8 months of age.
Article
Three classes of perceptual phenomena have repeatedly been associated with autism spectrum disorder (ASD): superior processing of fine detail (local structure), either inferior processing of overall/global structure or an ability to ignore disruptive global/contextual information, and impaired motion perception. This review evaluates the quality of the evidence bearing on these three phenomena. We argue that while superior local processing has been robustly demonstrated, conclusions about global processing cannot be definitively drawn from the experiments to date, which have generally not precluded observers using more local cues. Perception of moving stimuli is impaired in ASD, but explanations in terms of magnocellular/dorsal deficits do not appear to be sufficient. We suggest that abnormalities in the superior temporal sulcus (STS) may provide a neural basis for the range of motion-processing deficits observed in ASD, including biological motion perception. Such an explanation may also provide a link between perceptual abnormalities and specific deficits in social cognition associated with autism.