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Full-Text Articles in Behavior and Ethology

Dog And Owner Characteristics Predict Training Success, Jeffrey R. Stevens, London M. Wolff, Megan Bosworth, Jill Morstad Jan 2021

Dog And Owner Characteristics Predict Training Success, Jeffrey R. Stevens, London M. Wolff, Megan Bosworth, Jill Morstad

Jeffrey Stevens Publications

Teaching owners how to train their dogs is an important part of maintaining the health and safety of dogs and people. Yet we do not know what behavioral characteristics of dogs and their owners are relevant to dog training or if owner cognitive abilities play a role in training success. The aim of this study is to determine which characteristics of both dogs and owners predict success in completing the American Kennel Club Canine Good Citizen training program. Before the first session of a dog training course, owners completed surveys evaluating the behavior and cognition of their dog and themselves. …


Are Capuchin Monkeys (Sapajus Spp.) Sensitive To Lost Opportunities? The Role Of Opportunity Costs In Intertemporal Choice, Elsa Addessi, Valeria Tierno, Valentina Focaroli, Federica Rossi, Serena Gastaldi, Francesca De Petrillo, Fabio Paglieri, Jeffrey R. Stevens Jan 2021

Are Capuchin Monkeys (Sapajus Spp.) Sensitive To Lost Opportunities? The Role Of Opportunity Costs In Intertemporal Choice, Elsa Addessi, Valeria Tierno, Valentina Focaroli, Federica Rossi, Serena Gastaldi, Francesca De Petrillo, Fabio Paglieri, Jeffrey R. Stevens

Jeffrey Stevens Publications

Principles of economics predict that the costs associated with obtaining rewards can influence choice. When individuals face choices between a smaller, immediate option and a larger, later option, they often experience opportunity costs associated with waiting for delayed rewards because they must forego the opportunity to make other choices. We evaluated how reducing opportunity costs affects delay tolerance in capuchin monkeys. After choosing the larger option, in the High cost condition, subjects had to wait for the delay to expire, whereas in the Low cost different and Low cost same conditions, they could perform a new choice during the delay. …


Effects Of Human-Animal Interactions On Affect And Cognition, Elise L. Thayer, Jeffrey R. Stevens Jan 2021

Effects Of Human-Animal Interactions On Affect And Cognition, Elise L. Thayer, Jeffrey R. Stevens

Jeffrey Stevens Publications

Human-animal interaction has clear positive effects on people’s affect and stress. But less is known about how animal interactions influence cognition. We draw parallels between animal interactions and exposure to natural environments, a research area that shows clear improvements in cognitive performance. The aim of this study is to investigate whether interacting with animals similarly enhances cognitive performance, specifically executive functioning. To test this, we conducted two experiments in which we had participants self-report their affect and complete a series of cognitive tasks (long-term memory, attentional control, and working memory) before and after either a brief interaction with a dog …


Developing A Computer-Controlled Treat Dispenser For Canine Operant Conditioning, Walker Arce, Jeffrey R. Stevens Jan 2020

Developing A Computer-Controlled Treat Dispenser For Canine Operant Conditioning, Walker Arce, Jeffrey R. Stevens

Jeffrey Stevens Publications

When performing canine operant conditioning studies, the delivery of the reward can be a limiting factor of the study. While there are a few commercially available options for automatically delivering rewards, they generally require manual input, such as using a remote control, in accordance with the experiment script. This means that human reaction times and transmission distances can cause interruptions to the flow of the experiment. The potential for development of non-supervised conditioning studies is limited by this same factor. To remedy this, we retrofitted an off-the-shelf treat dispenser with new electronics that allow it to be remotely controllable as …


Mesotocin Influences Pinyon Jay Prosociality, Juan Duque, Whitney Leichner, Holly Ahmann, Jeffrey R. Stevens Jan 2018

Mesotocin Influences Pinyon Jay Prosociality, Juan Duque, Whitney Leichner, Holly Ahmann, Jeffrey R. Stevens

Jeffrey Stevens Publications

Many species exhibit prosocial behavior, in which one individual’s actions benefit another individual, often without an immediate benefit to itself. The neuropeptide oxytocin is an important hormonal mechanism influencing prosociality in mammals, but it is unclear whether the avian homologue mesotocin plays a similar functional role in birds. Here, we experimentally tested prosociality in pinyon jays (Gymnorhinus cyanocephalus), a highly social corvid species that spontaneously shares food with others. First, we measured prosocial preferences in a prosocial choice task with two different payoff distributions: Prosocial trials delivered food to both the subject and either an empty cage or a partner …


Replicability And Reproducibility In Comparative Psychology, Jeffrey R. Stevens May 2017

Replicability And Reproducibility In Comparative Psychology, Jeffrey R. Stevens

Jeffrey Stevens Publications

Psychology faces a replication crisis. The Reproducibility Project: Psychology sought to replicate the effects of 100 psychology studies. Though 97% of the original studies produced statistically significant results, only 36% of the replication studies did so (Open Science Collaboration, 2015). This inability to replicate previously published results, however, is not limited to psychology (Ioannidis, 2005). Replication projects in medicine (Prinz et al., 2011) and behavioral economics (Camerer et al., 2016) resulted in replication rates of 25 and 61%, respectively, and analyses in genetics (Munafò, 2009) and neuroscience (Button et al., 2013) question the validity of studies in those fields. Science, …


Review Of Bird Brain: An Exploration Of Avian Intelligence By Nathan Emery, Alan B. Bond Jan 2017

Review Of Bird Brain: An Exploration Of Avian Intelligence By Nathan Emery, Alan B. Bond

Avian Cognition Papers

Over the past 30 years, the study of bird behavior has been completely transformed by the ongoing revolution in cognitive psychology, opening up wholly new perspectives on the mental processes underlying such areas as foraging decisions, social intelligence, problem solving, memory encoding, and communication. Although these studies have contributed to a number of recent popular books, until now there has been no attempt to integrate avian cognition and recent findings in avian neuroanatomy and endocrinology into a single account that is attractive and accessible to a general readership. It is a steep challenge, but Nathan Emery has undertaken it in …


The Domain Specificity Of Intertemporal Choice In Pinyon Jays, Jeffrey R. Stevens, Bryce Kennedy, Dina Morales, Marianna Burks Jan 2016

The Domain Specificity Of Intertemporal Choice In Pinyon Jays, Jeffrey R. Stevens, Bryce Kennedy, Dina Morales, Marianna Burks

Jeffrey Stevens Publications

When choosing between a piece of cake now versus a slimmer waistline in the future, many of us have difficulty with self-control. Food-caching species, however, regularly hide food for later recovery, sometimes waiting months before retrieving their caches. It remains unclear whether these long-term choices generalize outside of the caching domain. We hypothesized that the ability to save for the future is a general tendency that cuts across different situations. To test this hypothesis, we measured and experimentally manipulated caching to evaluate its relationship with operant measures of self-control in pinyon jays (Gymnorhinus cyanocephalus). We found no correlation …


Reflections Of The Social Environment In Chimpanzee Memory: Applying Rational Analysis Beyond Humans, Jeffrey R. Stevens, Julian N. Marewski, Lael J. Schooler, Ian C. Gilby Jan 2016

Reflections Of The Social Environment In Chimpanzee Memory: Applying Rational Analysis Beyond Humans, Jeffrey R. Stevens, Julian N. Marewski, Lael J. Schooler, Ian C. Gilby

Jeffrey Stevens Publications

In cognitive science, the rational analysis framework allows modelling of how physical and social environments impose information-processing demands onto cognitive systems. In humans, for example, past social contact among individuals predicts their future contact with linear and power functions. These features of the human environment constrain the optimal way to remember information and probably shape how memory records are retained and retrieved. We offer a primer on how biologists can apply rational analysis to study animal behaviour. Using chimpanzees (Pan troglodytes) as a case study, we modelled 19 years of observational data on their social contact patterns. Much …


Voluntary Food Sharing In Pinyon Jays: The Role Of Reciprocity And Dominance, Juan F. Duque, Jeffrey R. Stevens Jan 2016

Voluntary Food Sharing In Pinyon Jays: The Role Of Reciprocity And Dominance, Juan F. Duque, Jeffrey R. Stevens

Jeffrey Stevens Publications

Food sharing offers a clear example of prosocial behavior, in which one individual’s actions benefit another. Researchers have proposed a range of hypotheses that explain why food sharing may occur among unrelated individuals. Two such hypotheses, reciprocity and dominance, have been tested in many species, including fish, corvids, rats, bats, and primates, showing that (1) recipients sometimes reciprocate sharing back to previous donors and (2) dominant individuals share more than subordinates. Although primates dominate the study of prosocial behavior, active donation of food is actually quite rare in primates. In contrast, several corvid species spontaneously share food much more frequently. …


Cognitive Phenotypes And The Evolution Of Animal Decisions, Tamra C. Mendelson, Courtney L. Fitzpatrick, Mark E. Hauber, Charles H. Pence, Rafael L. Rodriguez, Rebecca J. Safran, Caitlin A. Stern, Jeffrey R. Stevens Jan 2016

Cognitive Phenotypes And The Evolution Of Animal Decisions, Tamra C. Mendelson, Courtney L. Fitzpatrick, Mark E. Hauber, Charles H. Pence, Rafael L. Rodriguez, Rebecca J. Safran, Caitlin A. Stern, Jeffrey R. Stevens

Jeffrey Stevens Publications

Despite the clear fitness consequences of animal decisions, the science of animal decision making in evolutionary biology is underdeveloped compared with decision science in human psychology. Specifically, the field lacks a conceptual framework that defines and describes the relevant components of a decision, leading to imprecise language and concepts. The ‘judgment and decision-making’ (JDM) framework in human psychology is a powerful tool for framing and understanding human decisions, and we apply it here to components of animal decisions, which we refer to as ‘cognitive phenotypes’. We distinguish multiple cognitive phenotypes in the context of a JDM framework and highlight empirical …


Intertemporal Similarity: Discounting As A Last Resort, Jeffrey R. Stevens Jan 2016

Intertemporal Similarity: Discounting As A Last Resort, Jeffrey R. Stevens

Jeffrey Stevens Publications

Standard models of intertemporal choice assume that individuals discount future payoffs by integrating reward amounts and time delays to generate a discounted value. Alternative models propose that, rather than integrate across them, individuals compare within attributes (amounts and delays) to determine if differences in one attribute outweigh differences in another attribute. For instance, Leland (2002) and Rubinstein (2003) propose models that 1) compare the two reward amounts to determine whether they are similar, 2) compare the similarity of the two time delays, and then 3) make a decision based on these similarity judgments. Here, I tested discounting models against attribute-based …


Dangerous Mating Systems: Signal Complexity, Signal Content And Neural Capacity In Spiders, Marie E. Herberstein, Anne E. Wignall, Eileen Hebets, Jutta M. Schneider Oct 2014

Dangerous Mating Systems: Signal Complexity, Signal Content And Neural Capacity In Spiders, Marie E. Herberstein, Anne E. Wignall, Eileen Hebets, Jutta M. Schneider

Eileen Hebets Publications

Spiders are highly efficient predators in possession of exquisite sensory capacities for ambushing prey, combined with machinery for launching rapid and determined attacks. As a consequence, any sexually motivated approach carries a risk of ending up as prey rather than as a mate. Sexual selection has shaped courtship to effectively communicate the presence, identity, motivation and/or quality of potential mates, which help ameliorate these risks. Spiders communicate this information via several sensory channels, including mechanical (e.g. vibrational), visual and/or chemical, with examples of multimodal signaling beginning to emerge in the literature. The diverse environments that spiders inhabit have further shaped …


Changing Room Cues Reduces The Effects Of Proactive Interference In Clark’S Nutcrackers, Nucifraga Columbiana, Jody L. Lewis, Alan C. Kamil, Kate E. Webbink Jan 2013

Changing Room Cues Reduces The Effects Of Proactive Interference In Clark’S Nutcrackers, Nucifraga Columbiana, Jody L. Lewis, Alan C. Kamil, Kate E. Webbink

Avian Cognition Papers

To determine what factors are important for minimizing interference effects in spatial memory, Clark’s Nutcrackers, Nucifraga columbiana were tested for their spatial memory for two serial lists of locations per day. In this experiment two unique landmark sets were either different between List 1 and List 2 or the same. We found that Nutcrackers were most susceptible to interference when the landmark sets were the same. This study suggests that repeatedly testing animal memory in the same room, with the same cues, can hamper recall due to interference.


Differences In Relative Hippocampus Volume And Number Of Hippocampus Neurons Among Five Corvid Species, Kristy L. Gould, Karl E. Gilbertson, Andrew J. Hrvol, Joseph C. Nelson, Abigail L. Seyfer, Rose M. Brantner, Alan C. Kamil Jan 2013

Differences In Relative Hippocampus Volume And Number Of Hippocampus Neurons Among Five Corvid Species, Kristy L. Gould, Karl E. Gilbertson, Andrew J. Hrvol, Joseph C. Nelson, Abigail L. Seyfer, Rose M. Brantner, Alan C. Kamil

Avian Cognition Papers

The relative size of the avian hippocampus (Hp) has been shown to be related to spatial memory and food storing in two avian families, the parids and corvids. Basil et al. [Brain Behav Evol 1996;47: 156-164] examined North American food-storing birds in the corvid family and found that Clark’s nutcrackers had a larger relative Hp than pinyon jays and Western scrub jays. These results correlated with the nutcracker’s better performance on most spatial memory tasks and their strong reliance on stored food in the wild. However, Pravosudov and de Kort [Brain Behav Evol 67 (2006), 1-9] raised questions …


The Lives Of Others: Social Rationality In Animals, Jeffrey R. Stevens, Andrew J. King Jan 2013

The Lives Of Others: Social Rationality In Animals, Jeffrey R. Stevens, Andrew J. King

Jeffrey Stevens Publications

Darwin (1871) boldly claimed that humans and other animals differ very little in their cognition; a statement that has raised the ire of many scientists (Bolhuis & Wynne, 2009; Penn, Holyoak, & Povinelli, 2008). Rather than stating this continuity as a fact, we will explore the similarities and differences between humans and animals in the social rationality of their decision making. We find this a fruitful exercise because it can profit researchers of both humans and animals. Our purpose here is to provide an evolutionary background of social rationality: Why do animals attend to the lives of others? To this …


Do Clark’S Nutcrackers Demonstrate What-Where-When Memory On A Cache-Recovery Task?, Kristy L. Gould, Amy J. Ort, Alan C. Kamil Jan 2012

Do Clark’S Nutcrackers Demonstrate What-Where-When Memory On A Cache-Recovery Task?, Kristy L. Gould, Amy J. Ort, Alan C. Kamil

Avian Cognition Papers

What-where-when (WWW) memory during cache recovery was investigated in six Clark’s nutcrackers. During caching, both red- and blue-colored pine seeds were cached by the birds in holes filled with sand. Either a short (3 day) retention interval (RI) or a long (9 day) RI was followed by a recovery session during which caches were replaced with either a single seed or wooden bead depending upon the color of the cache and length of the retention interval. Knowledge of what was in the cache (seed or bead), where it was located, and when the cache had been made (3 or 9 …


Mechanisms For Decisions About The Future, Jeffrey R. Stevens Jan 2011

Mechanisms For Decisions About The Future, Jeffrey R. Stevens

Jeffrey Stevens Publications

Evolutionary and psychological perspectives on decision making remain largely separate endeavors. The bounded rationality approach integrates these two perspectives by focusing on simple, plausible mechanisms of decision making and the cognitive capacities needed to implement these mechanisms. Decisions about the future provide a class of decisions that lend themselves to a bounded rationality approach. Though many different mechanisms may exist for making decisions about the future, only a subset of these mechanisms actually require a representation of the future. The bounded rationality approach helps focus on the cognitive capacities and decision mechanisms that are necessary for a full understanding of …


Effects Of Gregariousness, Conspicuousness, And Novelty On Blue Jay (Cyanocitta Cristata) Learned Avoidance And Stimulus Generalization Of Unpalatable Prey, Joyce M. Dykema Apr 2008

Effects Of Gregariousness, Conspicuousness, And Novelty On Blue Jay (Cyanocitta Cristata) Learned Avoidance And Stimulus Generalization Of Unpalatable Prey, Joyce M. Dykema

Avian Cognition Papers

I examined a variety of factors hypothesized to be important in the evolution and maintenance of aposematism. Aposematism occurs when prey individuals advertise their toxic or otherwise aversive nature to potential predators via evolved conspicuous signals. I conducted three experiments in which blue jays (Cyanocitta cristata) were allowed to search a printed grayscale pixilated background for grayscale pixilated moths in an open room. I manipulated moth appearance and food reward, and recorded jay predation on the varying moth stimuli. In my first experiment, I repeated Alatalo & Mappes’ (1996) study examining the effects of prey gregariousness, or grouping, …


K-Means Clustering With Multiresolution Peak Detection, Guanshan Yu, Leen-Kiat Soh, Alan B. Bond May 2005

K-Means Clustering With Multiresolution Peak Detection, Guanshan Yu, Leen-Kiat Soh, Alan B. Bond

Avian Cognition Papers

Clustering is a practical data mining approach of pattern detection. Because of the sensitivity of initial conditions, k-means clustering often suffers from low clustering performance. We present a procedure to refine initial conditions of k-means clustering by analyzing density distributions of a data set before estimating the number of clusters k necessary for the data set, as well as the positions of the initial centroids of the clusters. We demonstrate that this approach indeed improves the accuracy and performance of k-means clustering measured by average intra to interclustering error ratio. This method is applied to the virtual ecology project to …


A Conceptual Framework For Non-Kin Food Sharing: Timing And Currency Of Benefits, Jeffrey R. Stevens, Ian C. Gilby Jan 2004

A Conceptual Framework For Non-Kin Food Sharing: Timing And Currency Of Benefits, Jeffrey R. Stevens, Ian C. Gilby

Jeffrey Stevens Publications

Many animal species, from arthropods to apes, share food. This paper presents a new framework that categorizes nonkin food sharing according to two axes: (1) the interval between sharing and receiving the benefits of sharing, and (2) the currency units in which benefits accrue to the sharer (especially food versus nonfood). Sharers can obtain immediate benefits from increased foraging efficiency, predation avoidance, mate provisioning, or manipulative mutualism. Reciprocity, trade, status enhancement and group augmentation can delay benefits. When benefits are delayed or when food is exchanged for nonfood benefits, maintaining sharing can become more difficult because animals face discounting and …


The Selfish Nature Of Generosity: Harassment And Food Sharing In Primates, Jeffrey R. Stevens Jan 2004

The Selfish Nature Of Generosity: Harassment And Food Sharing In Primates, Jeffrey R. Stevens

Jeffrey Stevens Publications

Animals may share food to gain immediate or delayed fitness benefits. Previous studies of sharing have concentrated on delayed benefits such as reciprocity, trade and punishment. This study tests an alternative model (the harassment or sharing–under–pressure hypothesis) in which a food owner immediately benefits because sharing avoids costly harassment from a beggar. I present an experiment that varies the potential ability of the beggar to harass, and of the owner to defend the food, to examine the effects of harassment on food sharing in two primate species: chimpanzees (Pan troglodytes) and squirrel monkeys (Saimiri boliviensis). For …


Discounting And Reciprocity In An Iterated Prisoner’S Dilemma, David W. Stephens, Colleen M. Mclinn, Jeffrey R. Stevens Jan 2002

Discounting And Reciprocity In An Iterated Prisoner’S Dilemma, David W. Stephens, Colleen M. Mclinn, Jeffrey R. Stevens

Jeffrey Stevens Publications

The Iterated Prisoner’s Dilemma (IPD) is a central paradigm in the study of animal cooperation. According to the IPD framework, repeated play (repetition) and reciprocity combine to maintain a cooperative equilibrium. However, experimental studies with animals suggest that cooperative behavior in IPDs is unstable, and some have suggested that strong preferences for immediate benefits (that is, temporal discounting) might explain the fragility of cooperative equilibria. We studied the effects of discounting and strategic reciprocity on cooperation in captive blue jays. Our results demonstrate an interaction between discounting and reciprocity. Blue jays show high stable levels of cooperation in treatments with …


The Effect Of Proximity On Landmark Use In Clark’S Nutcrackers, Kristy L. Gould-Beierle, Alan Kamil Jan 1999

The Effect Of Proximity On Landmark Use In Clark’S Nutcrackers, Kristy L. Gould-Beierle, Alan Kamil

Avian Cognition Papers

Clark’s nutcrackers, Nucifraga columbiana, store thousands of pine seeds during the autumn and retrieve them throughout the winter. It has been shown that these birds are able to use visual cues to relocate hidden food in the laboratory. In this set of experiments, we trained three groups of Clark’s nutcrackers to find a hidden food goal that was placed in the same spatial location relative to the testing room. During training, the location of two local cues in relation to the goal differed between the three groups. Group 1 learned the task with the cues closest to the goal, group …


Patterns Of Movement And Orientation During Caching And Recovery By Clark’S Nutcrackers, Nucifraga Columbiana, Alan Kamil, Russell P. Balda, Sally Good Jan 1999

Patterns Of Movement And Orientation During Caching And Recovery By Clark’S Nutcrackers, Nucifraga Columbiana, Alan Kamil, Russell P. Balda, Sally Good

Avian Cognition Papers

Clark’s nutcrackers regularly store large numbers of pine seeds and remember the locations of the cached seeds. Although they are very accurate, they do make some errors during recovery. In an attempt to determine whether any behaviours during caching predicted the occurrence of errors during recovery, we videotaped Clark’s nutcrackers while they cached and recovered seeds under laboratory conditions. We used the videotapes to develop complete, quantitative descriptions of caching and recovery behaviour, with an emphasis on body orientation and directions of movement. During caching, the birds showed the greatest change in their orientation and direction following cache creation. During …


How Do They, Indeed? A Reply To Biegler Et Al., Alan Kamil, Juli E. Jones Jan 1999

How Do They, Indeed? A Reply To Biegler Et Al., Alan Kamil, Juli E. Jones

Avian Cognition Papers

We trained Clark’s nutcrackers, Nucifraga columbiana, to search halfway between two landmarks while varying the distance between the landmarks (Kamil & Jones 1997). We found that the birds learned the problem readily and generalized to novel interlandmark distances within the range of distances used during training. Unlike some other studies in which responses to proportional distance were obtained (e.g. O’Keefe & Burgess 1996; Tommasi et al. 1997), the nutcrackers showed very precise search and maintained this precision during the transfer test. The distributions of digging locations around the central position were concentrated within ±1–2 cm of the central location …


The Ecology Of Foraging Behavior: Implications For Animal Learning And Memory, Alan Kamil, Herbert L. Roitblat Jan 1985

The Ecology Of Foraging Behavior: Implications For Animal Learning And Memory, Alan Kamil, Herbert L. Roitblat

Avian Cognition Papers

In his recent Annual Review of Psychology article, Snowdon (1983) discussed the synthesis of ethology and comparative psychology. A similar synthesis of behavioral ecology and animal learning is beginning to take place. This article reviews developments in the behavioral ecology and ethology of foraging behavior relevant to psychological research on animal learning. The psychological literature shows that animals possess a wide range of learning abilities, including “simple” classical and operant conditioning; they acquire spatial, nonspatial, and temporal discriminations; they exhibit various forms of rule learning (e.g. matching-to-sample and learning set), and may even in certain senses learn language. Why does …


Optimal Foraging Theory And The Psychology Of Learning, Alan Kamil Jan 1983

Optimal Foraging Theory And The Psychology Of Learning, Alan Kamil

Avian Cognition Papers

The development of optimization theory has made important contributions to the study of animal behavior. But the optimization approach needs to be integrated with other methods of ethology and psychology. For example, the ability to learn is an important component of efficient foraging behavior in many species, and the psychology of animal learning could contribute substantially to testing and extending the predictions of optimal foraging theory.


Spatial Learning As An Adaptation In Hummingbirds, Susan Cole, F. Reed Hainsworth, Alan Kamil, Terre Mercier, Larry L. Wolf Jan 1982

Spatial Learning As An Adaptation In Hummingbirds, Susan Cole, F. Reed Hainsworth, Alan Kamil, Terre Mercier, Larry L. Wolf

Avian Cognition Papers

An ecological approach based on food distribution suggests that hummingbirds should more easily learn to visit a flower in a new location than to learn to return to a flower in a position just visited, for a food reward. Experimental results support this hypothesis as well as the general view that differences in learning within and among species represent adaptations.


Visual Detection Of Cryptic Prey By Blue Jays (Cyanocitta Cristata), Alexandra T. Pietrewicz, Alan Kamil Jan 1977

Visual Detection Of Cryptic Prey By Blue Jays (Cyanocitta Cristata), Alexandra T. Pietrewicz, Alan Kamil

Avian Cognition Papers

Blue jays learned to respond differentially to the presence or absence of Catocala moths in slides. This detection of the moths by the jays was affected by the background upon which the moth was placed and its body orientation, thus providing an objective measure of crypticity. These procedures are useful for the study of visual detection of prey.