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"The angular interval between the direction of progression and body orientation in normal, alcohol- and cocaine treated fruit flies."
Anna Gakamsky, Efrat Oron, Dan Valente, Partha P Mitra, Daniel Segal, Yoav Benjamini and Ilan Golani (2013)
PloS one. Vol. 8 (10) , pp. e76257.
Abstract: In this study we characterize the coordination between the direction a fruit-fly walks and the direction it faces, as well as offer a methodology for isolating and validating key variables with which we phenotype fly locomotor behavior. Our fundamental finding is that the angular interval between the direction a fly walks and the direction it faces is actively managed in intact animals and modulated in a patterned way with drugs. This interval is small in intact flies, larger with alcohol and much larger with cocaine. The dynamics of this interval generates six coordinative modes that flow smoothly into each other. Under alcohol and much more so under cocaine, straight path modes dwindle and modes involving rotation proliferate. To obtain these results we perform high content analysis of video-tracked open field locomotor behavior. Presently there is a gap between the quality of descriptions of insect behaviors that unfold in circumscribed situations, and descriptions that unfold in extended time and space. While the first describe the coordination between low-level kinematic variables, the second quantify cumulative measures and subjectively defined behavior patterns. Here we reduce this gap by phenotyping extended locomotor behavior in terms of the coordination between low-level kinematic variables, which we quantify, combining into a single field two disparate fields, that of high content phenotyping and that of locomotor coordination. This will allow the study of the genes/brain/locomotor coordination interface in genetically engineered and pharmacologically manipulated animal models of human diseases.
Annotation: In this paper fruit fly behavior is characterized (phenotyped) in terms of the dynamics of the coordination between the two degrees of freedom available to the fly at the scale of the animal's path – the animal's shift of weight (direction of translation of the animal's center) and the animal's front (the direction the fly is facing). The two drug preparations are used as two phenotypes whose dynamics is compared to the dynamics of intact fly behavior.
"Short and long term measures of anxiety exhibit opposite results."
Ehud Fonio, Yoav Benjamini and Ilan Golani (2012)
PloS one. Vol. 7 (10) , pp. e48414.
Abstract: Animal models of human diseases of the central nervous system, generalized anxiety disorder included, are essential for the study of the brain-behavior interface and obligatory for drug development; yet, these models fail to yield new insights and efficacious drugs. By increasing testing duration hundredfold and arena size tenfold, and comparing the behavior of the common animal model to that of wild mice, we raise concerns that chronic anxiety might have been measured at the wrong time, for the wrong duration, and in the wrong animal. Furthermore, the mice start the experimental session with a short period of transient adaptation to the novel environment (habituation period) and a long period reflecting the respective trait of the mice. Using common measures of anxiety reveals that mice exhibit opposite results during these periods suggesting that chronic anxiety should be measured during the post-habituation period. We recommend tools for measuring the transient period, and provide suggestions for characterizing the post habituation period.
Annotation: This is a twin article, to be read before the "Nature Methods" article "Measuring behavior of animal models: faults and remedies".
"Measuring behavior of animal models: faults and remedies."
Ehud Fonio, Ilan Golani and Yoav Benjamini (2012)
Nature methods. Vol. 9 (12) , pp. 1167-70.
Abstract: Widely used behavioral assays need re-evaluation and validation against their intended use. We focus here on measures of chronic anxiety in mouse models and posit that widely used assays such as the open-field test are performed at the wrong time, for inadequate durations and using inappropriate mouse strains. We propose that behavioral assays be screened for usefulness on the basis of their replicability across laboratories.
Annotation: The absence of a wide perspective in time, space and natural history, and the use of ad hoc measures having no intrinsic (Statistical and/or geometrical) justification yield animal models that have poor translational values for human diseases of the brain.
"The developmental dynamics of behavioral growth processes in rodent egocentric and allocentric space."
Ilan Golani (2012)
Behavioural brain research. Vol. 231 (2) , pp. 309-16.
Abstract: In this review I focus on how three methodological principles advocated by Philip Teitelbaum influenced my work to this day: that similar principles of organization should be looked for in ontogeny and recovery of function; that the order of emergence of behavioral components provides a view on the organization of that behavior; and that the components of behavior should be exhibited by the animal itself in relatively pure form. I start by showing how these principles influenced our common work on the developmental dynamics of rodent egocentric space, and then proceed to describe how these principles affected my work with Yoav Benjamini and others on the developmental dynamics of rodent allocentric space. We analyze issues traditionally addressed by physiological psychologists with methods borrowed from ethology, EW (Eshkol-Wachman) movement notation, dynamical systems and exploratory data analysis. Then we show how the natural origins of axes embodied by the behavior of the organism itself, are used by us as the origins of axes for the measurement of the developmental moment-by-moment dynamics of behavior. Using this methodology we expose similar principles of organization across situations, species and preparations, provide a developmental view on the organization of behavior, expose the natural components of behavior in relatively pure form, and reveal how low level primitives generate higher level constructs. Advances in tracking technology should allow us to study how movements in egocentric and allocentric spaces interlace. Tracking of multi-limb coordination, progress in online recording of neural activity in freely moving animals, and the unprecedented accumulation of genetically engineered mouse preparations makes the behavioral ground plan exposed in this review essential for a systematic study of the brain/behavior interface.
Annotation: We assume that organisms do not move in a haphazard way. Therefore  we first search for the origin established by the organism itself for performing a particular behavior.  Then we follow the organism step by step, measuring its behavior in reference to that origin. This procedure exposes the intrinsic dimensionality, the building blocks, and the developmental dynamics of the behavior. Here, we expose the phenomenology (a la Husserl) of mice experiencing their egocentric and allocentric spaces.
"Validation of the dimensionality emergence assay for the measurement of innate anxiety in laboratory mice."
Apar Jain, Anna Dvorkin, Ehud Fonio, Ilan Golani and Cornelius T Gross (2012)
European neuropsychopharmacology : the journal of the European College of Neuropsychopharmacology. Vol. 22 (2) , pp. 153-63.
Abstract: The open field test is a common tool to measure innate anxiety in rodents. In the usual configuration of this test the animal is forced to explore the open arena and its behavior includes both anxiety and non-anxiety responses. However, the open arena is generally small and allows only limited expression of exploratory behavior. The recently developed dimensionality emergence assay in which an animal is housed in a home cage with free access to a large circular arena elicits graded exploration and promises to serve as a more ethological test of anxiety. Here we examined the predictive validity of this assay for anxiety-related measures in mice. First, we compared their behavior in the presence or absence of access to the home cage and found that mice with access to the home cage exhibited a gradual build-up in exploration of the arena while those without did not. Then we identified behavioral measures that responded to treatment with the anxiolytic drug diazepam. Diazepam altered several classical measures of innate anxiety, such as distance traveled and thigmotaxis, but also led to a dose-dependent acceleration of the build-up as reflected in a significantly reduced latency to attain several exploratory landmarks. Finally, we tested the utility of the dimensionality emergence assay in assessing alterations in innate anxiety reported in mice carrying a knockout allele for the serotonin 1A receptor (Htr1a). Our findings support the validity of the dimensionality emergence assay as a method to extract an expanded repertoire of behavioral measures for the assessment of anxiety in laboratory mice.
"Quantifying the buildup in extent and complexity of free exploration in mice"
Yoav Benjamini, Ehud Fonio, Tal Galili, Gregor Z Havkin and Ilan Golani (2011)
Proceedings of the National Academy of Sciences of the United States of America. Vol. 108 Suppl , pp. 15580-7.
Abstract: To obtain a perspective on an animal's own functional world, we study its behavior in situations that allow the animal to regulate the growth rate of its behavior and provide us with the opportunity to quantify its moment-by-moment developmental dynamics. Thus, we are able to show that mouse exploratory behavior consists of sequences of repeated motion: iterative processes that increase in extent and complexity, whose presumed function is a systematic active management of input acquired during the exploration of a novel environment. We use this study to demonstrate our approach to quantifying behavior: targeting aspects of behavior that are shown to be actively managed by the animal, and using measures that are discriminative across strains and treatments and replicable across laboratories.
"Longitudinal Assessment of Deliberate Mouse Behavior in the Home Cage and Attached Environments: Relevance to Anxiety and Mood Disorders"
MJ Kas, I Golani and Y Benjamini (2011)
, In Mood and Anxiety Related Phenotypes in Mice. Vol. 63 , pp. 1--20.
Abstract: Understanding behavioral regulation can further progress by developing new approaches that allow refinement of behavioral phenotypes. The current availability of several thousand different mutant mice and of human candidate genes for emotional (affective) disorders challenges behavioral neuroscientists to extend their views and methodologies to dissect complex behaviors into behavioral phenotypes and subsequently to define gene–behavioral phenotype relationships. Here, we put forward multiday automated behavioral and physiological observations in carefully designed environments to assess evolutionary conserved behavioral strategies in mice. This offers the opportunity to design experimental setups that allow the animals themselves to regulate their own behavior, using representations of continuous kinematic variables, studying the dynamics of behavior (change across time or change across activity); i.e., growth or decay processes of behavior and concomitant physiological adjustments such as heart rate. The measures characterizing these processes should have discriminative power (across strains or treatments) and be replicable (across laboratories). Furthermore, cross species genetic studies for these neurobehavioral and physiological traits may provide a novel way toward identifying neurobiological mechanisms underlying core features of complex psychiatric disorders.
"Ten ways to improve the quality of descriptions of whole-animal movement."
Yoav Benjamini, Dina Lipkind, Guy Horev, Ehud Fonio, Neri Kafkafi and Ilan Golani (2010)
Neuroscience and biobehavioral reviews. Vol. 34 (8) , pp. 1351-65.
Abstract: The demand for replicability of behavioral results across laboratories is viewed as a burden in behavior genetics. We demonstrate how it can become an asset offering a quantitative criterion that guides the design of better ways to describe behavior. Passing the high benchmark dictated by the replicability demand requires less stressful and less restraining experimental setups, less noisy data, individually customized cutoff points between the building blocks of movement, and less variable yet discriminative dynamic representations that would capture more faithfully the nature of the behavior, unmasking similarities and differences and revealing novel animal-centered measures. Here we review ten tools that enhance replicability without compromising discrimination. While we demonstrate the usefulness of these tools in the context of inbred mouse exploratory behavior they can readily be used in any study involving a high-resolution analysis of spatial behavior. Viewing replicability as a design concept and using the ten methodological improvements may prove useful in many fields not necessarily related to spatial behavior.
"Knots: attractive places with high path tortuosity in mouse open field exploration."
Anna Dvorkin, Henry Szechtman and Ilan Golani (2010)
PLoS computational biology. Vol. 6 (1) , pp. e1000638.
Abstract: When introduced into a novel environment, mammals establish in it a preferred place marked by the highest number of visits and highest cumulative time spent in it. Examination of exploratory behavior in reference to this "home base" highlights important features of its organization. It might therefore be fruitful to search for other types of marked places in mouse exploratory behavior and examine their influence on overall behavior.Examination of path curvatures of mice exploring a large empty arena revealed the presence of circumscribed locales marked by the performance of tortuous paths full of twists and turns. We term these places knots, and the behavior performed in them-knot-scribbling. There is typically no more than one knot per session; it has distinct boundaries and it is maintained both within and across sessions. Knots are mostly situated in the place of introduction into the arena, here away from walls. Knots are not characterized by the features of a home base, except for a high speed during inbound and a low speed during outbound paths. The establishment of knots is enhanced by injecting the mouse with saline and placing it in an exposed portion of the arena, suggesting that stress and the arousal associated with it consolidate a long-term contingency between a particular locale and knot-scribbling.In an environment devoid of proximal cues mice mark a locale associated with arousal by twisting and turning in it. This creates a self-generated, often centrally located landmark. The tortuosity of the path traced during the behavior implies almost concurrent multiple views of the environment. Knot-scribbling could therefore function as a way to obtain an overview of the entire environment, allowing re-calibration of the mouse's locale map and compass directions. The rich vestibular input generated by scribbling could improve the interpretation of the visual scene.
"High-throughput data analysis in behavior genetics"
Anat Sakov, Ilan Golani, Dina Lipkind and Yoav Benjamini (2010)
The Annals of Applied Statistics. Vol. 4 (2) , pp. 743-763.
Abstract: In recent years, a growing need has arisen in different fields for the development of computational systems for automated analysis of large amounts of data (high-throughput). Dealing with nonstandard noise structure and outliers, that could have been detected and corrected in manual analysis, must now be built into the system with the aid of robust methods. We discuss such problems and present insights and solutions in the context of behavior genetics, where data consists of a time series of locations of a mouse in a circular arena. In order to estimate the location, velocity and acceleration of the mouse, and identify stops, we use a nonstandard mix of robust and resistant methods: LOWESS and repeated running median. In addition, we argue that protection against small deviations from experimental protocols can be handled automatically using statistical methods. In our case, it is of biological interest to measure a rodent’s distance from the arena’s wall, but this measure is corrupted if the arena is not a perfect circle, as required in the protocol. The problem is addressed by estimating robustly the actual boundary of the arena and its center using a nonparametric regression quantile of the behavioral data, with the aid of a fast algorithm developed for that purpose.
"Freedom of movement and the stability of its unfolding in free exploration of mice"
Ehud Fonio, Yoav Benjamini and Ilan Golani (2009)
Proceedings of the National Academy of Sciences of the United States of America. Vol. 106 (50) , pp. 21335-40.
Abstract: Exploration is a central component of human and animal behavior that has been studied in rodents for almost a century. The measures used by neuroscientists to characterize full-blown exploration are limited in exposing the dynamics of the exploratory process, leaving the morphogenesis of its structure and meaning hidden. By unfettering exploration from constraints imposed by hunger, thirst, coercion, and the confines of small cage and short session, using advanced computational tools, we reveal its meaning in the operational world of the mouse. Exploration consists of reiterated roundtrips of increasing amplitude and freedom, involving an increase in the number of independent dimensions along which the mouse moves (macro degrees of freedom). This measurable gradient can serve as a standard reference scale for the developmental dynamics of some aspects of the mouse's emotional-cognitive state and for the study of the interface between behavior and the neurophysiologic and genetic processes mediating it.
"Mouse cognition-related behavior in the open-field: emergence of places of attraction."
Anna Dvorkin, Yoav Benjamini and Ilan Golani (2008)
PLoS computational biology. Vol. 4 (2) , pp. e1000027.
Abstract: Spatial memory is often studied in the Morris Water Maze, where the animal's spatial orientation has been shown to be mainly shaped by distal visual cues. Cognition-related behavior has also been described along "well-trodden paths"--spatial habits established by animals in the wild and in captivity reflecting a form of spatial memory. In the present study we combine the study of Open Field behavior with the study of behavior on well-trodden paths, revealing a form of locational memory that appears to correlate with spatial memory. The tracked path of the mouse is used to examine the dynamics of visiting behavior to locations. A visit is defined as either progressing through a location or stopping there, where progressing and stopping are computationally defined. We then estimate the probability of stopping at a location as a function of the number of previous visits to that location, i.e., we measure the effect of visiting history to a location on stopping in it. This can be regarded as an estimate of the familiarity of the mouse with locations. The recently wild-derived inbred strain CZECHII shows the highest effect of visiting history on stopping, C57 inbred mice show a lower effect, and DBA mice show no effect. We employ a rarely used, bottom-to-top computational approach, starting from simple kinematics of movement and gradually building our way up until we end with (emergent) locational memory. The effect of visiting history to a location on stopping in it can be regarded as an estimate of the familiarity of the mouse with locations, implying memory of these locations. We show that the magnitude of this estimate is strain-specific, implying a genetic influence. The dynamics of this process reveal that locations along the mouse's trodden path gradually become places of attraction, where the mouse stops habitually.
"Coordination of steering in a free-trotting quadruped"
Eyal Gruntman, Yoav Benjamini and Ilan Golani (2007)
Journal of comparative physiology. A, Neuroethology, sensory, neural, and behavioral physiology. Vol. 193 (3) , pp. 331-45.
Abstract: Typically, locomotion has been studied by restricting the animal's path and/or speed, focusing on stride and step kinematics. Here we incorporate measurements of the legs and trunk in the support and swing phases, during trotting with various speeds and curvatures. This paradigm releases the animal from the confines of the treadmill and runway into the open space. The diagonal step, a new unit of locomotion, is defined by regarding the line between the two supporting diagonal legs as a frame of reference for the description of the dynamics of the virtual line between the two swinging diagonal legs. This analysis reveals that during free trotting the mouse uses three types of steps: fixating, opening, and closing steps. During progression along a straight path, the mouse uses fixating steps, in which the swinging diagonal maintains a fixed direction, landing on the supporting foreleg; during progression along a curved path the mouse uses opening and closing steps alternately. If two steps of the same type are performed sequentially, they engender an abrupt change of direction. Our results reveal how steering with the swinging diagonal, while using a virtually bipedal gait, engenders the whole repertoire of free-trotting behavior.
"Estimating wall guidance and attraction in mouse free locomotor behavior."
G Horev, Y Benjamini, A Sakov and I Golani (2007)
Genes, brain, and behavior. Vol. 6 (1) , pp. 30-41.
Abstract: In this study, we estimate the influence exerted by the wall of the Open Field on the trajectory of the mouse. The wall exerts two types of influence on the mouse's path: one of guidance and one of attraction. The guiding influence is expressed by the tendency of mice to progress in parallel to the wall. This tendency wanes with increasing distance from the wall but is observed at large distances from it. The more parallel the mouse is to the wall the higher is its speed, even when distant from the wall. This association between heading direction and speed shows that the mouse controls its heading in reference to the wall. It is also observed in some blind strains, revealing that wall-guidance is not based exclusively on vision. The attraction influence is reflected by movement along the wall and by the asymmetry between speed during movement toward, and during movement away from the wall: sighted mice move faster toward the wall, whereas blind mice use similar speeds in both directions. Measures characterizing these influences are presented for five inbred strains, revealing heritable components that are replicable across laboratories. The revealed structure can lead to the identification of distinct groups of genes that mediate the distinct influences of guidance and attraction exerted by the wall. It can also serve as a framework for the decoding of electrophysiological data recorded in free moving rodents in the Open Field.
"Analysis of the trajectory of Drosophila melanogaster in a circular open field arena."
Dan Valente, Ilan Golani and Partha P Mitra (2007)
PloS one. Vol. 2 (10) , pp. e1083.
Abstract: BACKGROUND: Obtaining a complete phenotypic characterization of a freely moving organism is a difficult task, yet such a description is desired in many neuroethological studies. Many metrics currently used in the literature to describe locomotor and exploratory behavior are typically based on average quantities or subjectively chosen spatial and temporal thresholds. All of these measures are relatively coarse-grained in the time domain. It is advantageous, however, to employ metrics based on the entire trajectory that an organism takes while exploring its environment.

METHODOLOGY/PRINCIPAL FINDINGS: To characterize the locomotor behavior of Drosophila melanogaster, we used a video tracking system to record the trajectory of a single fly walking in a circular open field arena. The fly was tracked for two hours. Here, we present techniques with which to analyze the motion of the fly in this paradigm, and we discuss the methods of calculation. The measures we introduce are based on spatial and temporal probability distributions and utilize the entire time-series trajectory of the fly, thus emphasizing the dynamic nature of locomotor behavior. Marginal and joint probability distributions of speed, position, segment duration, path curvature, and reorientation angle are examined and related to the observed behavior.

CONCLUSIONS/SIGNIFICANCE: The measures discussed in this paper provide a detailed profile of the behavior of a single fly and highlight the interaction of the fly with the environment. Such measures may serve as useful tools in any behavioral study in which the movement of a fly is an important variable and can be incorporated easily into many setups, facilitating high-throughput phenotypic characterization.

"Characterizing Animal Behavior through Audio and Video Signal Processing"
Dan Valente, Haibin Wang, Peter Andrews, Partha P. Mitra, Sigal Saar, Ofer Tchernichovski, Ilan Golani and Yoav Benjamini (2007)
IEEE Multimedia. Vol. 14 (4) , pp. 32-41.
Abstract: This article presents two instances in which multimedia systems and processing have elucidated animal behavior and have been central in developing quantitative descriptions. These examples demonstrate multimedia systems' utility and necessity in developing a complete phenotypic description. We hope that this article will spur interest in this subject in the multimedia community, so more advanced processing techniques will enter the field of quantitative neuroethology. You might have noticed that in our two examples, there was nothing very multimodal about the media techniques used. Both of these systems are transparently unimodal. This speaks to the limited crossover between the multimedia community and the behavioral neuroscientists (or neuroethologists). These examples did show, however, that the neuroscientific community can benefit greatly from incorporating multimedia techniques into their experiments and data analysis. As the walls between these disciplines begin to fall, experimental setups that are truly multimedia will likely appear. Such systems will allow complete phenotypic descriptions of animals in ethologically relevant settings, along with methods for analyzing, manipulating, annotating, and storing the resulting data. Combining these phenotypic descriptions with the corresponding genetic and neural network properties will facilitate the connection of these organization levels and lead to a more thorough understanding of brain functioning.
"Multimedia signal processing for behavioral quantification in neuroscience"
Peter Andrews, Haibin Wang, Dan Valente, Jihène Serkhane, Partha P. Mitra, Sigal Saar, Ofer Tchernichovski and Ilan Golani (2006)
, In Proceedings of the 14th annual ACM international conference on Multimedia - MULTIMEDIA '06. , pp. 1007.
Abstract: While there have been great advances in quantification of the genotype of organisms, including full genomes for many species, the quantification of phenotype is at a comparatively primitive stage. Part of the reason is technical difficulty: the phenotype covers a wide range of characteristics, ranging from static morphological features, to dynamic behavior. The latter poses challenges that are in the area of multimedia signal processing. Automated analysis of video and audio recordings of animal and human behavior is a growing area of research, ranging from the behavioral phenotyping of genetically modified mice or drosophila to the study of song learning in birds and speech acquisition in human infants. This paper reviews recent advances and identifies key problems for a range of behavior experiments that use audio and video recording. This research area offers both research challenges and an application domain for advanced multimedia signal processing. There are a number of MMSP tools that now exist which are directly relevant for behavioral quantification, such as speech recognition, video analysis and more recently, wired and wireless sensor networks for surveillance. The research challenge is to adapt these tools and to develop new ones required for studying human and animal behavior in a high throughput manner while minimizing human intervention. In contrast with consumer applications, in the research arena there is less of a penalty for computational complexity, so that algorithmic quality can be maximized through the utilization of larger computational resources that are available to the biomedical researcher.
"Wild mouse open field behavior is embedded within the multidimensional data space spanned by laboratory inbred strains."
E Fonio, Y Benjamini, A Sakov and I Golani (2006)
Genes, brain, and behavior. Vol. 5 (5) , pp. 380-8.
Abstract: The vast majority of studies on mouse behavior are performed on laboratory mouse strains (Mus laboratorius), while studies of wild-mouse behavior are relatively rare. An interesting question is the relationship between the phenotypes of M. laboratorius and the phenotypes of their wild ancestors. It is commonly believed, often in the absence of hard evidence, that the behavior of wild mice exceeds by far, in terms of repertoire richness, magnitude of variables and variability of behavioral measures, the behavior of the classical inbred strains. Having phenotyped the open field behavior (OF) of eight of the commonly used laboratory inbred strains, two wild-derived strains and a group of first-generation-in-captivity local wild mice (Mus musculus domesticus), we show that contrary to common belief, wild-mouse OF behavior is moderate, both in terms of end-point values and in terms of their variability, being embedded within the multidimensional data space spanned by laboratory inbred strains. The implication could be that whereas natural selection favors moderate locomotor behavior in wild mice, the inbreeding process tends to generate in mice, in some of the features, extreme and more variable behavior.
"Activity density in the open field: a measure for differentiating the effect of psychostimulants."
Neri Kafkafi and Gregory I. Elmer (2005)
Pharmacology, biochemistry, and behavior. Vol. 80 (2) , pp. 239-49.
Abstract: Traditional open-field activity measures do not provide a sharp behavioral differentiation across psychomotor stimulants such as d-amphetamine (AMPH) and cocaine (COC) in the mouse. We used Software for the Exploration of Exploration (SEE) to investigate and develop a novel behavioral endpoint to characterize the "structure" of AMPH- and COC-induced locomotor behavior in two inbred strains of mouse, C57BL/6 (B6) and DBA/2 (D2). We suggest a measure we term "activity density" as a means to differentiate the behavioral effects of COC and AMPH. Activity density is defined as the activity divided by the range over which it took place. It characterizes the restriction of behavioral repertoire that does not result merely from inactivity. In both the B6 and D2 mice, AMPH increased activity density in a dose-dependent fashion by restricting the range of activity compared with COC doses producing the same level of activity. While AMPH restricted the range in both genotypes, characterizing the geographical region in which the restriction took place further differentiated the genotypes. The newly developed activity density measure thus provides a more general measure than stereotypy of the path, and can differentiate the effects of AMPH and COC both within and across genotypes.
"Genotype-environment interactions in mouse behavior: a way out of the problem."
Neri Kafkafi, Yoav Benjamini, Anat Sakov, Greg I Elmer and Ilan Golani (2005)
Proceedings of the National Academy of Sciences of the United States of America. Vol. 102 (12) , pp. 4619-24.
Abstract: In behavior genetics, behavioral patterns of mouse genotypes, such as inbred strains, crosses, and knockouts, are characterized and compared to associate them with particular gene loci. Such genotype differences, however, are usually established in single-laboratory experiments, and questions have been raised regarding the replicability of the results in other laboratories. A recent multilaboratory experiment found significant laboratory effects and genotype x laboratory interactions even after rigorous standardization, raising the concern that results are idiosyncratic to a particular laboratory. This finding may be regarded by some critics as a serious shortcoming in behavior genetics. A different strategy is offered here: (i) recognize that even after investing much effort in identifying and eliminating causes for laboratory differences, genotype x laboratory interaction is an unavoidable fact of life. (ii) Incorporate this understanding into the statistical analysis of multilaboratory experiments using the mixed model. Such a statistical approach sets a higher benchmark for finding significant genotype differences. (iii) Develop behavioral assays and endpoints that are able to discriminate genetic differences even over the background of the interaction. (iv) Use the publicly available multilaboratory results in single-laboratory experiments. We use software-based strategy for exploring exploration (see) to analyze the open-field behavior in eight genotypes across three laboratories. Our results demonstrate that replicable behavioral measures can be practically established. Even though we address the replicability problem in behavioral genetics, our strategy is also applicable in other areas where concern about replicability has been raised.
"Texture of locomotor path: a replicable characterization of a complex behavioral phenotype."
N Kafkafi and G I Elmer (2005)
Genes, brain, and behavior. Vol. 4 (7) , pp. 431-43.
Abstract: A database of mouse locomotor path in spatial tests can be used to search in silico for behavioral measures that better discriminate between genotypes and are more replicable across laboratories. In this study, software for the exploration of exploration (SEE) was used to search a large database for a novel behavioral measure that would characterize complex movement paths. The database included mouse open-field behavior assessed in 3 laboratories, 7 inbred strains, several pharmacological treatments and hundreds of animals. The new behavioral measure, "path texture", was characterized using the local curvature of the path (the change of direction per unit distance, in degrees/cm) across several spatial scales, starting from scales smaller than the animal's body length and up to the scale of the arena size. Path texture analysis differs from fractal dimension analysis in that it does not assume self-similarity across scales. Path texture was found to discriminate inbred strains with relatively high broad-sense heritability (4371 and high replicability across laboratories. Even genotypes that had similar path curvatures in some scales usually differed in other scales, and self-similarity across scales was not displayed by all genotypes. Amphetamine decreased the path curvature of C57BL/6 mice in small and medium scales, while having no effect on DBA/2J mice. Diazepam dose-dependently decreased the curvature of C57BL/6 mice across all scales, while 2 anxiogenic drugs, FG-7142 and pentylenetetrazole, increased it. Path texture thus has high potential for behavioral phenotyping and the study of drug effects in the mouse.
"The Behavior of the Laboratory Rat : A Handbook with Tests: A Handbook with Tests"
Ilan Golani, Yoav Benjamini, A Dvorkin, Dina Lipkind and Neri Kafkafi (2004)
, pp. 520.
Abstract: More is known about the behavior, anatomy, and molecular biology of the laboratory rat than any other animal species. Although its natural history and psychological functions have been described previously in books, this is the first comprehensive description of its behavior. Both seasoned and beginning investigators will be amazed at the range and complexity of the species as described in the 43 chapters of this volume. The behavioral descriptions are closely tied to the laboratory methods from which they were derived, thus allowing investigators to correlate the behavior and methods and exploit them in their own research. This book is aimed at investigators in neuroscience who may not be familiar with rat behavior, but who wish to incorporate behavioral studies into their own research. Nevertheless, seasoned investigators will also find the book to be a handy reference for behavioral paradigms with which they may not be familiar. It is expected that as the genetic and molecular understanding of the rat develops, there will be an increasing need for knowledge about rat behavior. This book will serve as an indispensable resource for neuroscientists, psychologists, pharmacologists, geneticists, molecular biologists, zoologists, and their students and trainees.
"The dynamics of spatial behavior: how can robust smoothing techniques help?"
Itay Hen, Anat Sakov, Neri Kafkafi, Ilan Golani and Yoav Benjamini (2004)
Journal of Neuroscience Methods. Vol. 133 (1-2) , pp. 161-172.
Abstract: A variety of setups and paradigms are used in the neurosciences for automatically tracking the location of an animal in an experiment and for extracting features of interest out of it. Many of these features, however, are critically sensitive to the unavoidable noise and artifacts of tracking. Here, we examine the relevant properties of several smoothing methods and suggest a combination of methods for retrieving locations and velocities and recognizing arrests from time series of coordinates of an animal’s center of gravity. We accomplish these by using robust nonparametric methods, such as Running Median (RM) and locally weighted regression methods. The smoothed data may, subsequently, be segmented to obtain discrete behavioral units with proven ethological relevance. New parameters such as the length, duration, maximal speed, and acceleration of these units provide a wealth of measures for, e.g., mouse behavioral phenotyping, studies on spatial orientation in vertebrates and invertebrates, and studies on rodent hippocampal function. This methodology may have implications for many tests of spatial behavior.
"New replicable anxiety-related measures of wall vs center behavior of mice in the open field."
Dina Lipkind, Anat Sakov, Neri Kafkafi, Gregory I Elmer, Yoav Benjamini and Ilan Golani (2004)
Journal of applied physiology (Bethesda, Md. : 1985). Vol. 97 (1) , pp. 347-59.
Abstract: Anxiety is a widely studied psychiatric disorder and is thought to be a complex and multidimensional phenomenon. Sensitive behavioral discrimination of animal models of anxiety is crucial for the elucidation of the behavioral components of anxiety and the physiological processes that mediate them. Commonly used behavior paradigms of anxiety usually include only a few automatically collected measures; these do not exhaust the behavioral richness exhibited by animals, thus perhaps missing important differences between preparations. The aim of the present study was to expand the repertoire of automatically collected measures in a classical test of anxiety: behavior in relation to the wall in the open field. We present an algorithm, based on the Software for the Exploration of Exploration strategy, which automatically partitions the mouse path into intrinsically defined patterns of movement near the wall and in the center. These patterns are used to design new end points, which provide an articulated description of various aspects of behavior near the wall and in the center. Sixteen new end points were designed with data from C57BL/6J and DBA/2J mice tested in three laboratories. The strain differences in all end points were evaluated on another data set to assess their validity and were found to remain stable. Ten of the sixteen end points were found to discriminate between the two strains in a replicable manner. The entire set of end points can be used on various genetic and pharmacological models of anxiety with good prospects of providing fine discrimination in a replicable manner.
"Extending SEE for large-scale phenotyping of mouse open-field behavior"
Neri Kafkafi (2003)
Behavior Research Methods, Instruments, & Computers. Vol. 35 (2) , pp. 294-301.
Abstract: SEE (Software for the Exploration of Exploration) is a visualization and analysis tool designed for the study of open-field behavior in rodents. In this paper, I present new extensions of SEE that were designed to facilitate its use for mouse behavioral phenotyping and, especially, for the problems of discrimination of genotypes and the replication of results across laboratories and experimental conditions. These extensions were specifically designed to promote a new approach in behavioral phenotyping, reminiscent of the approach that has been successfully employed in bioinformatics during recent years. The path coordinates of all animals from many experiments are stored in a database. SEE can be used to query, visualize, and analyze any desirable subsection of this database and to design new measures (endpoints) with increasingly better discriminative power and replicability. The use of the new extensions is demonstrated here in the analysis of results from several experiments and laboratories, with an emphasis on this approach.
"Darting behavior: a quantitative movement pattern designed for discrimination and replicability in mouse locomotor behavior"
Neri Kafkafi, Michal Pagis, Dina Lipkind, Cheryl L Mayo, Yoav Bemjamini, Ilan Golani and Gregory I Elmer (2003)
Behavioural brain research. Vol. 142 (1-2) , pp. 193-205.
Abstract: In the open-field behavior of rodents, Software for Exploring Exploration (SEE) can be used for an explicit design of behavioral endpoints with high genotype discrimination and replicability across laboratories. This ability is demonstrated here in the development of a measure for darting behavior. The behavior of two common mouse inbred strains, C57BL/6J (B6) and DBA/2J (D2), was analyzed across three different laboratories, and under the effect of cocaine or amphetamine. "Darting" was defined as having higher acceleration during progression segments while moving less during stops. D2 mice darted significantly more than B6 mice in each laboratory, despite being significantly less active. These differences were maintained following cocaine administration (up to 20mg/kg) and only slightly altered by amphetamine (up to 5mg/kg) despite a several fold increase in activity. The replicability of darting behavior was confirmed in additional experiments distinct from those used for its design. The strategy leading to the darting measure may be used to develop additional discriminative and replicable endpoints of open-field behavior.
"SEE locomotor behavior test discriminates C57BL/6J and DBA/2J mouse inbred strains across laboratories and protocol conditions."
Neri Kafkafi, Dina Lipkind, Yoav Benjamini, Cheryl L Mayo, Gregory I Elmer and Ilan Golani (2003)
Behavioral neuroscience. Vol. 117 (3) , pp. 464-77.
Abstract: Conventional tests of behavioral phenotyping frequently have difficulties differentiating certain genotypes and replicating these differences across laboratories and protocol conditions. This study explores the hypothesis that automated tests can be designed to quantify ethologically relevant behavior patterns that more readily characterize heritable and replicable phenotypes. It used SEE (Strategy for the Exploration of Exploration) to phenotype the locomotor behavior of the C57BL/6 and DBA/2 mouse inbred strains across 3 laboratories. The 2 genotypes differed in 15 different measures of behavior, none of which had a significant genotype-laboratory interaction. Within the same laboratory, most of these differences were replicated in additional experiments despite the test photoperiod phase being changed and saline being injected. Results suggest that well-designed tests may considerably enhance replicability across laboratories.
"Controlling the false discovery rate in behavior genetics research"
Yoav Benjamini, Dan Drai, Greg Elmer, Neri Kafkafi and Ilan Golani (2001)
Behavioural Brain Research. Vol. 125 (1-2) , pp. 279-284.
Abstract: The screening of many endpoints when comparing groups from different strains, searching for some statistically significant difference, raises the multiple comparisons problem in its most severe form. Using the 0.05 level to decide which of the many endpoints' differences are statistically significant, the probability of finding a difference to be significant even though it is not real increases far beyond 0.05. The traditional approach to this problem has been to control the probability of making even one such error--the Bonferroni procedure being the most familiar procedure achieving such control. However, the incurred loss of power stemming from such control led many practitioners to neglect multiplicity control altogether. The False Discovery Rate (FDR), suggested by Benjamini and Hochberg [J Royal Stat Soc Ser B 57 (1995) 289], is a new, different, and compromising point of view regarding the error in multiple comparisons. The FDR is the expected proportion of false discoveries among the discoveries, and controlling the FDR goes a long way towards controlling the increased error from multiplicity while losing less in the ability to discover real differences. In this paper we demonstrate the problem in two studies: the study of exploratory behavior [Behav Brain Res (2001)], and the study of the interaction of strain differences with laboratory environment [Science 284 (1999) 1670]. We explain the FDR criterion, and present two simple procedures that control the FDR. We demonstrate their increased power when used in the above two studies.
"Rats and mice share common ethologically relevant parameters of exploratory behavior."
D Drai, N Kafkafi, Y Benjamini, G Elmer and I Golani (2001)
Behavioural brain research. Vol. 125 (1-2) , pp. 133-40.
Abstract: Detailed studies of rat exploratory behavior reveal that it consists of typical behavior patterns having a distinct structure. Recently we have developed interactive software that uses as input the automatically digitized time-series of the animal's location for the visualization, analysis, capturing and quantification of these patterns. We use this software here for the study of BALB/cJtau mouse behavior. The results suggest that a considerable number of rat patterns are also present in the mouse. These ethologically-relevant patterns have a significant potential as a phenotyping tool.
"SEE: a tool for the visualization and analysis of rodent exploratory behavior"
Dan Drai and Ilan Golani (2001)
Neuroscience & Biobehavioral Reviews. Vol. 25 (5) , pp. 409-426.
Abstract: The complexity of exploratory behavior creates a need for a visualization and analysis tool that will highlight regularities and help generating new hypotheses about the structure of this behavior. The hypotheses can then be formulated as algorithms that capture the patterns and quantify them. SEE is a Mathematica based software developed by us for the exploration of exploratory behavior. The raw data for SEE are a time series of the animal ‘s coordinates in space sampled at a rate that allows a meaningful computation of speeds. SEE permits: (i) a visualization of the path of the animal and a computation of the dynamics of activity; (ii) a decomposition of the path into several modes of motion (1st gear, 2nd gear, etc.) and a computation of the typical maximal speeds, the spatial spread, and the proportion of each of these modes; and(iii) a visualization of the location in the environment of stopping episodes, along with their dwell time. These visualizations highlight the presence of preferred places, including the animal's so-called home base, and permits a computation of the spatio-temporal diversity in the location of stopping episodes. The software also: (i) decomposes the animal's path into round trips from the home base, called ‘excursions’, and computes the number of stops per excursion; (ii) generates a visualization of the phase space (path+speed, traced in a three-dimensional graph) of any progression segment or list of such segments; and (iii) produces a visualization of the way places in the animal's operational world are connected to each other. SEE also permits the definition and computation of behavioral endpoints across any section of any database of raw data. The range of applicability of SEE to various experimental setups, tracking procedures, species, and preparations is addressed in the discussion.
"Natural segmentation of the locomotor behavior of drug-induced rats in a photobeam cage"
N Kafkafi, C Mayo, D Drai, I Golani and G Elmer (2001)
Journal of neuroscience ldots. Vol. 109 (2) , pp. 111-21.
Abstract: Recently, Drai et al. (J Neurosci Methods 96 (2000) 119) have introduced an algorithm that segments rodent locomotor behavior into natural units of 'staying in place' (lingering) behavior versus going between places (progression segments). This categorization, based on the maximum speed attained within the segment, was shown to be intrinsic to the data, using the statistical method of Gaussian Mixture Model. These results were obtained in normal rats and mice using very large (650 or 320 cm) circular arenas and a video tracking system. In the present study, we reproduce these results with amphetamine, phencyclidine and saline injected rats, using data measured by a standard photobeam tracking system in square 45 cm cages. An intrinsic distinction between two or three 'gears' could be shown in all animals. The spatial distribution of these gears indicates that, as in the large arena behavior, they correspond to the difference between 'staying in place' behavior and 'going between places'. The robustness of this segmentation over arena size, different measurement system and dose of two psychostimulant drugs indicates that this is an intrinsic, natural segmentation of rodent locomotor behavior. Analysis of photobeam data that is based on this segmentation has thus a potential use in psychopharmacology research.
"Statistical discrimination of natural modes of motion in rat exploratory behavior."
D Drai, Y Benjamini and I Golani (2000)
Journal of neuroscience methods. Vol. 96 (2) , pp. 119-31.
Abstract: We analyze the locomotor behavior of the rat during exploration, and show that digitally collected data (time series of positions) provide a sufficient basis for establishing that the rat uses several distinct modes of motion (first, second, third, and sometimes fourth gear). The distinction between these modes is obtained by first segmenting the time series into sequences of data points occurring between arrests (as ascertained within the resolution of the data acquisition system). The statistical distribution of the maximal amount of motion occurring within each of these episodes is then analyzed and shown to be multi modal. This enables us to decompose motion into distinct modes. In one application of this decomposition we show that the ethological ad hoc notion of stopping behavior corresponds to progression without leaving first gear. We do so by showing that the spatial spread of such progressions is confined to a small 20-50 cm range in a 6.5 m diameter arena. This provides a justification for a construct of 'staying in place'. This construct is not defined in terms of position in objective space, but purely in terms of the rat's own behavior. We test the generality of our method by applying it to mouse exploratory behavior.
"Phenotyping stereotypic behaviour: collective variables, range of variation and predictability"
I Golani, N Kafkafi and D Drai (1999)
Applied Animal Behaviour Science. Vol. 65 (3) , pp. 191-220.
Abstract: Stereotypy is the narrowing down of an animal's behavioural repertoire. Starting from normal behaviour there is a continuum between rich and free behaviour on the one hand, and dull and predictable stereotypies on the other. To study the continuum all that one needs are “knobs” (like drugs or stress) that shift the behaviour from one end to the other, and a method for documenting the behavioural shift. The degree of narrowing down of the animal's repertoire is measurable in terms of the number of “collective variables” available to the animal, the range of values each collective variable can take, and the predictability of sequences of movements. We substantiate this thesis by presenting examples taken from the normal and drug-induced behaviour of the rat.
"A traveling wave of lateral movement coordinates both turning and forward walking in the ferret"
Neri Kafkafi and Ilan Golani (1998)
Biological cybernetics. Vol. 78 (6) , pp. 441-453.
Abstract: Relative phase was recently suggested as a key variable for the dynamical modeling of coordination in both quadruped locomotion and undulation swimming in fish. Relative phase analysis has not yet been applied, however, to the behavior of intact, freely moving animals, but only to simplified situations involving restrained animals and humans. In order to investigate relative phase under free movement conditions, we filmed free locomotion of ferrets (Mustella putorius) from below (through a glass floor) and measured the lateral bending along the head, torso, and tail, and the location of the four paws. We introduced an algorithm which extracts the phase (and thus also the relative phase) even when the movements were neither periodic nor symmetric. Our results show that relative phases between segments have preferred values, which are relatively independent of the amplitude, duration, and asymmetry of the movement. In particular, both walking and turning can be explained as modulations of a single pattern: a cephalo-caudal, traveling wave of lateral movement with a wavelength of approximately one length of the body. The relative phase between movements of adjacent segments is similar when the body is in S shape (i.e., when walking forward), or C shape (i.e., when turning). The movements of the paws in the horizontal plane can also be considered as part of this traveling wave. Our findings suggest that the concept of traveling waves of lateral bending, as found in the locomotion of undulating fish, can be generalized in two ways: (i) by considering the axis around which the movement is centered, it applies not only to forward locomotion, but also to turning; (ii) by incorporating the position of the paws, it applies also to the movement of quadrupeds. Our findings suggest that the relative phase, once it is generalized to asymmetric and quasi-periodic movement, is suitable for modeling coordination patterns under free movement conditions.
"The dynamics of long-term exploration in the rat. Part I. A phase-plane analysis of the relationship between location and velocity."
O Tchernichovski, Y Benjamini and I Golani (1998)
Biological cybernetics. Vol. 78 (6) , pp. 423-32.
Abstract: Rat exploratory behavior consists of regular excursions into the environment from a preferred place termed a home base. A phase plane representation of excursions reveals a geometrical pattern that changes during exploration in both shape and size. We first show that with time and repeated exposures to the same large environment there is a gradual increase in the length of excursions; each rat has its own characteristic length of excursions; but all rats share a similar rate of excursion growth. As in our experimental setup the rats perform increasingly longer paths from one location, while locomoting back and forth along the walls of the arena, exposure is more extensive at the proximal part of the route, and less at the distal part. We consequently show that the rat's velocity pattern changes concurrently with the increase in excursion length, and in correlation with the level of exposure (familiarity) to places. The primitive velocity pattern consists of slow progression while moving away from base and fast progression while returning to it. During exposure the asymmetry in velocity is inverted. The inversion spreads across successive excursions from the home base outwards. The rate of spread of this inversion is higher than the rate of increase in excursion length, and is similar across rats. Because it spreads more rapidly than the increase in excursion length, the global shape of the excursion trajectory changes. The dynamics of excursion shape share similar properties with the dynamics of excursion length. Both might reflect the same intrinsic constraints on the amount of novelty that a rat can handle per excursion.
"The dynamics of long-term exploration in the rat. Part II. An analytical model of the kinematic structure of rat exploratory behavior."
O Tchernichovski and Y Benjamini (1998)
Biological cybernetics. Vol. 78 (6) , pp. 433-40.
Abstract: A simple analytical model is proposed here that captures to a large extent the kinematic structure of rat exploratory behavior. Previous studies have shown that such behavior consists of regular excursions into the environment from a preferred place termed a home base. In the first part of this study, we showed that with time and repeated exposure to the same large environment, there is a gradual increase in the length of excursions. Concurrently, the rat's velocity pattern changes in a complex yet structured way, which is correlated with the exposure (= familiarity) to places. In this part, we show that the complex pattern described there might be explained by an analytic model, in terms of a simple dynamical system, with few assumptions concerning motivation and learning. The model is studied both by analysis and simulation. The theoretical examination of the dynamics of excursion length suggests that excursion length increases as a linear function of two system parameters, one governing the rate of motivation loss, and the other the rate of (location-specific) familiarization. Combining this theoretical finding with the empirical results suggests that the two theoretical parameters are linearly related: the less confident the rat, the slower its familiarization rate, and thus differences in patterns of movement between rats can be explained using one rat-specific parameter. Furthermore, the more complex velocity pattern of the rat can then be easily captured by the same model. The analyzed behavior of the rat suggests that the locale sensory information that the rat collects has a gradient towards the home base, with decreasing information input away from home base. This sensory pattern emerges from the simple set of rules and restrictions on the rat's exploratory behavior. Thus, instead of imposing a set of ad hoc restrictions on a simulated rat so that its spatial learning is similar to that of a real rat, the model suggests a set of simple intrinsic constraints to govern the exploratory behavior.
"Keeping the Body Straight in the Unconstrained Locomotion of Normal and Dopamine-Stimulant-Treated Rats."
I. Golani, H. Einat, O. Tchernichovski and P. Teitelbaum (1997)
Journal of motor behavior. Vol. 29 (2) , pp. 99-112.
Abstract: During unconstrained locomotor behavior, rats move in and out of a straight posture of the body (including the head). In the present study, the stability of maintaining a straight body was examined in untreated rats and in rats treated with saline (SAL) or with I of 3 dopamine stimulants (n = 4 rats per group). The stability of maintaining a straight body can range from very high (with 0.5 mg/kg quinpirole [QUIN], to high (first half-session with 5 mg/kg (+)-amphetamine [AMPHD], to very low (second half-session with 5 mg/kg AMPH), or can be maintained at a level similar to that observed in untreated rats (with 1.25mg/kg apomorphine [APO].Stability was assessed by videotaping the rats and, then, by using frame-by-frame analysis, scoring the cumulative proportion of time spent in a straight posture, the frequency of transitions from one hemisphere to the other without being trapped in the midline plane, and the degree of lateral bending during turning and during walking on a curved path. The present study is one in a series identifying key variables that constrain as many degrees of freedom as possible in rat locomotor behavior. The uncovering of such variables is an indispensible step that precedes dynamic systems stability analysis and provides candidates for key variables for the modeling of motor coordination
"Coordination of side-to-side head movements and walking in amphetamine-treated rats: a stereotyped motor pattern as a stable equilibrium in a dynamical system"
Neri Kafkafi, Stavit Levi-Havusha, Ilan Golani and Yoav Benjamini (1996)
Biological Cybernetics. Vol. 495 (6) , pp. 487-495.
Abstract: Rats injected with 5.0 mg/kg (+)-amphetamine perform, at one stage of the drug's influence, rhythmic side-to-side head movements while walking. This makes them an interesting preparation for investigating how stereotyped motor patterns emerge from the coordination of periodic movements. We report here such a pattern we have isolated: the left foreleg and the right hindleg land on the ground as the head reaches the peak of its movement to the right, and vice versa (contra-lateral pattern). We show that this pattern can be explained as a stable equilibrium in a simple, nonlinear dynamical model, similar to models developed for tapping with both hands in human subjects. The model also accounts for sequences of behavior that are not in the contra-lateral pattern, explaining them as a flow of the system back towards the stable equilibrium after a disturbance. Motor patterns that constitute the building blocks of unconstrained behavior are often defined as fixed phase relations between movements of the parts of the body. This study applies the paradigm of Dynamic Pattern Generation to free (unconstrained) behavior: motor patterns are defined as stable equilibria in dynamical systems, assembled by mutual influence of concurrent movements. Our findings suggest that this definition is more powerful for the description of free behavior. The amphetamine-treated rat is a useful preparation for investigating this notion in an unconstrained animal whose behavior is still not as complex and variable as that of the normal animal
"Constraints and the Emergence of 'Free' Exploratory Behavior in Rat Ontogeny"
Ofer Tchernichovski, Yoav Benjamini and Ilan Golani (1996)
Behaviour. Vol. 133 (7) , pp. 519-539.
Abstract: The present study attempts to combine the study of spatial learning with the study of open field behavior. We examine rat moment-to-moment behavior in the wide context of i) a large testing environment, ii) repeated exposures, and iii) development. Previous studies have shown that in adult rats, exploratory behavior of a novel environment is organized around a reference place termed the rat's home base. In this study we show that the appearance of a homebase is a singular stage in ontogeny, marking the transition from a low to a high scatter of movement in the environment. The increase in scatter is characterized by the appearance of several additional reference places. We suggest that the rat connects these reference places gradually and in a regular fashion. To do so we employ statistical filters which extract the principal places visited by the rat, and use measures of diversity which estimate the scatter of movement around these places. The presented data are the first derived from unconstrained behavior, supporting the hypothesis that the rat's cognitive space is represented in terms of local charts eventually combined into a global map.
"A phase plane representation of rat exploratory behavior."
O Tchernichovski and I Golani (1995)
Journal of neuroscience methods. Vol. 62 (1-2) , pp. 21-7.
Abstract: Rat spontaneous spatial behavior is considered to be stochastic and is therefore commonly analyzed in terms of cumulative measures. Here, we suggest a method which generates a moment-to-moment representation of this behavior. It has been proposed earlier that rat spatial behavior can be partitioned into natural units termed excursions (round trips) performed from a reference place termed the rat's home base. We offer a phase plane representation of excursions (plotting the rat's momentary location against its momentary velocity). The results reveal a geometrical pattern, typical of young age and early exposure. It consists of low velocity and intermittent progression while moving away from the home base (upstream segment), and high velocity while moving back to it (downstream segment). The asymmetry between the two segments defines a field of significance in the rat's operational world. This field undergoes regular transformations, revealing thereby the rat's strategy of occupancy of the environment. The presented dynamics could provide a framework for the interpretation of concurrent neural events associated with navigation and spatial memory.
"The practicality of using the Eshkol-Wachman movement notation in behavioral pharmacology and kinesics"
Ilan Golani (1994)
Behavioral and Brain Sciences. Vol. 17 (04) , pp. 754.
"Stopping behavior: constraints on exploration in rats (Rattus norvegicus)."
I Golani, Y Benjamini and D Eilam (1993)
Behavioural brain research. Vol. 53 (1-2) , pp. 21-33.
Abstract: In the absence of an obvious reference place, rat locomotor behavior in a novel environment appears haphazard. In previous work, one or two places termed home bases, were shown to stand out from all the other places in the environment in terms of the behaviors performed in them and in terms of their behavioral stability. We use home base location as a reference place for rat movement in locale space, by defining an excursion as a trip starting at a home base and ending at the next stop at a home base. We then establish the uniform distribution as an appropriate model for the number of stops per excursion. This way we show that there is an intrinsic upper bound on the number of times a rat stops during an excursion. As a rat leaves the home base, home base attraction increases with every additional stop performed by it, first slowly and then fast. This cumulative process of attraction may be concluded after each stop, as long as the number of stops does not exceed an intrinsic upper bound; once the upper bound is reached, the rat concludes that excursion and returns to base. The session's upper bound does not increase with the size of the explored area.
"Tail display in Atractaspis engaddensis (Astractaspididae, Serpentes)"
E. Kochva and I. Golani (1993)
Copeia. (1) , pp. 226-228.
"Oscillators in the human body and circular-muscle gymnastics."
S Yom-Tov and I Golani (1993)
Medical hypotheses. Vol. 41 (2) , pp. 118-22.
Abstract: There is a growing body of literature about the role of oscillators in the living body, and about the interactions between different oscillators. Considering the importance of endogenous oscillators in regulating the body's functions, and the existence of 'dynamical diseases', diseases of control systems which involve oscillators in the body, a way to mend dysfunctioning oscillators seems to be needed. Circular-muscle gymnastics, a method of physical activity which has been developed in Israel, reveals some phenomena which may point in a promising direction. Some of these phenomena call to mind known facts and theories about oscillators and their effects.
"The mobility gradient as an integrating model in the organization of vertebrate mowement"
Ilan Golani (1992)
Behavioral and Brain Sciences. Vol. 15 (02) , pp. 249-266.
Abstract: Ordinary language can prevent us from seeing the organization of whole-animal movement. This may be why the search for behavioral homologies has not been as fruitful as the founders of ethology had hoped. The Eshkol-Wachman (EW) movement notational system can reveal shared movement patterns that are undetectable in the kinds of informal verbal descriptions of the same behaviors that are in current use. Rules of organization that are common to locomotor development, agonistic and exploratory behavior, scent marking, play, and dopaminergic drug-induced stereotypies in a variety of vertebrates suggest that behavior progresses along a “mobility gradient” from immobility to increasing complexity and unpredictability. A progression in the opposite direction, with decreasing spatial complexity and increased stereotypy, occurs under the influence of the nonselective dopaminergic drugs apomorphine and amphetamine and partly also the selective dopamine agonist quinpirole. The behaviors associated with the mobility gradient appear to be mediated by a family of basal ganglia-thalamocortical circuits and their descending output stations. Because the small number of rules underlying the mobility gradient account for a large variety of behaviors, they may be related to the specific functional demands on these neurological systems. The EW system and the mobility gradient model should prove useful to ethologists and neurobiologists.
"The description of rat drug-induced behavior: kinematics versus response categories."
N Adani, N Kiryati and I Golani (1991)
Neuroscience and biobehavioral reviews. Vol. 15 (4) , pp. 455-60.
Abstract: The study of rat drug-induced locomotor behavior is largely based on the assumption that behavior consists of a sequence of response categories performed by the whole animal one at a time. By analysing this behavior under (+)-amphetamine (5 mg/kg), we illustrate how even a precise definition of such categories may not be sufficient for the establishment of behavioral variables that have a "physiological reality." We describe the changes of relation between the parts of the rat's body in reference to selected coordinate systems, and show that a great variety of locomotor patterns observed under amphetamine can be reduced to as few as 3 descriptive component-variables. These continuous and relatively independent variables, which behave predictably in the course of drug action, operate simultaneously. Variations in their relative timing of onset and termination account for the apparent variability in observed behavior. The economy and generality of a description based on these variables suggest the existence of corresponding central mechanisms of control.
"Home base behavior in amphetamine-treated tame wild rats (Rattus norvegicus)"
D. Eilam and I. Golani (1990)
Behavioural Brain Research. Vol. 36 (1-2) , pp. 161-170.
Abstract: When a rat treated with amphetamine (0.5-5 mg/kg) locomotes in an unfamiliar environment, there are one or two places which it visits most often. In these one or two places the mean duration of a visit (stop) is the longest, and, compared to other places, the incidence of grooming and rearing are the highest. Since in intact rats these features of place characterize it as a 'home base', it is concluded that under amphetamine rats also establish one or two home bases. One home base was generally established by rats treated with low doses of amphetamine, while two bases were most evident in those treated with high doses. Since the paths of locomotion in amphetamine-treated rats were previously described to be stereotyped, it is suggested that home base location under this drug may be used as a reference point in the assessment of the organization of stereotyped locomotor behavior.
"Evidence that apomorphine and (+)-amphetamine produce different types of circling in rats."
a R Cools, W Scheenen, D Eilam and I Golani (1989)
Behavioural brain research. Vol. 34 (1-2) , pp. 111-6.
Abstract: Apomorphine and (+)-amphetamine are known to produce circling in naive rats. Frame by frame analysis of videotape recordings of the behaviour of Wistar rats treated with a subcutaneous injection of apomorphine (1.1 mg/kg; n = 8) or (+)-amphetamine (0.5 and 1.0 mg/kg; n = 8 and n = 8) was used to study this behaviour in more detail. In line with previously reported studies, apomorphine was found to change the functioning of hindlimb stepping. In contrast, (+)-amphetamine was found to change the functioning of forelimb stepping. These data imply that apomorphine and (+)-amphetamine produce their drug-specific circling via different substrates within the brain.
"Home base behavior of rats (Rattus norvegicus) exploring a novel environment."
D. Eilam and I. Golani (1989)
Behavioural brain research. Vol. 34 (3) , pp. 199-211.
Abstract: When rats are placed in a novel environment, they alternate between progression and stopping: in the course of a session they stop briefly in many places, but in one or two places they also stop for very long periods. The place in which they stay for the longest cumulative time is defined as the rat's home base. In this place the incidences of grooming and of rearing are high and often the highest. In addition, the number of visits to the home base is typically the highest. Some rats establish a secondary base with similar properties to those of the main home base. The location of the base influences the mode of progression throughout the environment: progression away from base is slower and includes more stops than progression back. It is suggested that this paradigm may be used for the analysis of the spatial organization of locomotor behavior in neuroscience research.
"D2-agonist quinpirole induces perseveration of routes and hyperactivity but no perseveration of movements"
David Eilam, Ilan Golani and Henry Szechtman (1989)
Brain Research. Vol. 490 (2) , pp. 255-267.
Abstract: The behavior in an open field of rats injected with the D2-agonist quinpirole (2 mg/kg; n = 10) and saline (n = 10) was analyzed in terms of routes and movements. Quinpirole induces perserveration of routes without inducing perseveration of movements. Perseveration of routes consists of repeated travel along a few paths in a limited portion of the environment. Lack of perseveration of movements was evidenced by the same distribution of lateral, vertical, and forward movements as in saline-treated animals. Quinpirole also increased the total amount of progression and the total number of movements performed by the rat's body parts along all dimensions of movements. Thus, under quinpirole, animals were hyperactive, stereotyped in route, but free in movement. This profile resembles behavior under low doses of amphetamine but not the behavior under either apomorphine or high doses of amphetamine. Thus, contrary to the current view, administration of a D2-receptor agonist is sufficient to produce a major component of dopamine-induced stereotyped behavior. It is suggested that quinpirole induces perseveration of route by affecting presynaptic release of dopamine, and that the organization of route is independent of the organization of movement.
"The ontogeny of exploratory behavior in the house rat (Rattus rattus): the mobility gradient"
D Eilam and I Golani (1988)
Developmental psychobiology. Vol. 21 (7) , pp. 679-710.
Abstract: Infants of rats and other mammals respond to a novel environment by becoming immobile, and then showing a process of motorial expansion called "warm-up." Starting from immobility, new types of movement are incorporated into the stream of behavior according to rather strict rules of order. Once a new type of movement has been performed, the infant reverts to it repeatedly. As a result, the earlier portion of the behavior appears stereotyped, giving the impression of an automatism. Later, as new types of movement are added to the infant's repertoire, the movement becomes increasingly rich and unpredictable, giving the impression of "free" behavior. The same rules of order operate within "warm-up" sequences of movement, and across such sequences, day by day. Concurrently, there is an increase in the amplitude of movements, resulting in a gradual expansion of the portion of the environment explored by the infant. The same rules of order seem to operate in the development of locomotion in more primitive vertebrates. In rats under the action of psychoactive drugs, the "warm-up" sequence is performed in reverse.
"Striking and other offensive and defensive behavior patterns in Atractaspis engaddensis (Ophidia, Atractaspididae)"
I. Golani and E. Kochva (1988)
Copeia. (3) , pp. 792-797.
"A different look at measurement and interpretation of drug-induced stereotyped behavior"
Henry Szechtman, David Eilam, Philip Teitelbaum and Ilan Golani (1988)
Psychobiology. Vol. 16 (2) , pp. 164-173.
"Superiority and Inferiority: a Morphological Analysis of Free and Stimulus Bound Behaviour in Honey Badger (Mellivora capensis) Interactions"
YONA YANIV and Ilan Golani (1987)
Ethology. Vol. 74 (2) , pp. 89-116.
Abstract: In analyzing motor behaviour, we use a method of gestalt perception that relies less on intuition and more on rational processes. The elementary building blocks of behaviour in this study are single movements — distinct changes of relation between two adjacent parts of the body. The single movements are performed in groups around specific axes: whole body rotations around the vertical absolute axis, whole body rotations around the longitudinal axis of the body, and whole body rotations around the side-to-side axis of the body. In their full blown form these rotations amount, respectively, to pivoting, rolling and tumbling. During “ritualized fighting”, these rotations are incorporated into the behaviour of the badger in a fixed sequence, yielding a fugue of rotations. “Ritualized fighting” is comprised in the badgers of five higher level (whole body) building blocks: the three whole body rotations, squatting, and forward walking. These building blocks are termed in the present study component-variables. At a still higher level of analysis we reveal the effect of the environment (the moving partner), on the performance of these component-variables. We record continuously the parts of the bodies of the two partners that touch or almost touch each other. In this way, we specify the tactile and visual input which impinges on each of the partners at any one time in the course of the interaction. Then, we examine the effect of the very same input, on the types of response performed by the badgers. This method allows us to assess the freedom of movement, i.e. the number of different responses available to each of the partners when confronted with the same stimulus situation. The five component-variables generate four composite profiles of actual behaviour (the “inferior” female, the “inferior” male, the “superior” female, and the “superior” male). They form a common denominator in all the profiles, but vary systematically in amount, amplitude, and frequency from one profile to the next, yielding a gradient. Part of the gradient was also described in “ritualized fighting” in wolves, and in other species and situations. In wolves and badgers it involves a gradual transition from relative immobility in the most inferior, to extensive mobility in the most superior partner. We show, that the same stimulus situations elicit in the inferior the most fixed response sequences, and in the superior, the most variable ones. Inferiority consists of relative immobility and stimulus bound behaviour; superiority consists of extensive mobility and relatively free behaviour. The difference between fixed response sequences and “voluntary” behaviour is of degree, not of principle.
"Adaptability of innate motor patterns and motor control mechanisms"
M. B. Berkinblit, A. G. Feldman and O. I. Fukson (1986)
Behavioral and Brain Sciences. Vol. 9 (04) , pp. 585.
Abstract: The following factors underlying behavioral plasticity are discussed: (1) reflex adaptability and its role in the voluntary control of movement, (2) degrees of freedom and motor equivalence, and (3) the problem of the discrete organization of motor behavior. Our discussion concerns a variety of innate motor patterns, with emphasis on the wiping reflex in the frog. It is proposed that central regulation of stretch reflex thresholds governs voluntary control over muscle force and length. This suggestion is an integral part of the equilibrium-point hypothesis, two versions of which are compared. Kinematic analysis of the wiping reflex in the spinal frog has shown that each stimulated skin site is associated with a group of different but equally effective trajectories directed to the target site. Such phenomena reflect the principle of motor equivalence -the capacity of the neuronal structures responsible for movement to select one or another of a set of possible trajectories leading to the goal. Redundancy of degrees of freedom at the neuronal level as well as at the mechanical level of the body's joints makes motor equivalence possible. This sort of equivalence accommodates the overall flexibility of motor behavior. An integrated behavioral act or a single movement consists of dynamic components. We distinguish six components for the wiping reflex, each associated with a certain functional goal, specific body positions, and motor-equivalent movement patterns. The nervous system can combine the available components in various ways in forming integrated behavioral sequences. The significance of command neuronal organization is discussed with respect to (1) the combinatory strategy of the nervous system and (2) the relation between continuous and discrete forms of motor control. We conclude that voluntary movements are effected by the central nervous system with the help of the mechanisms that underlie the variability and modifiability of innate motor patterns.
"What are the building blocks of the frog's wiping reflex?"
Ilan Golani (1986)
, In Behavioral and Brain Sciences. Vol. 9 (04) , pp. 607.
Abstract: Examines some of the descriptive tools used by M. B. Berkinblit et al (see PA, Vol 75:9852) for the representation and analysis of movement in their study of innate motor patterns and motor control mechanisms. ((c) 1997 APA/PsycINFO, all rights reserved)
"Early ontogeny of face grooming in mice."
I Golani and J C Fentress (1985)
Developmental psychobiology. Vol. 18 (6) , pp. 529-44.
Abstract: A U-shaped function of behavioral growth characterizes the early ontogeny of face grooming in mice. In the first few postnatal days (0-100 hr), mice groom their face by using temporally isolated strokes, or bouts of strokes, which vary in amplitude and symmetry. Later on (100-200 hr), bouts disappear, asymmetry is eliminated, and the amplitude of strokes is gradually restricted; the infants engage in stereotyped, double-handed, temporally isolated strokes. Finally (200-300 hr), bouts reappear, including both short and long, symmetrical and asymmetrical, strokes. These changes, are accompanied by unidirectional changes such as an increasing participation of the trunk, the neck, and the head in grooming, which lead to the flexible organization of face grooming seen in adults. Instead of describing development in terms of emerging unitary "acts," we have recorded from high-speed films three simultaneous aspects of movement: the movements of the separate limb and body segments, the resultant paths which are traced by the forepaws, and the paths of contact which are traced on the face. This method of description discloses (a) reversible changes, and (b) a change in the relative stability of each of these aspects of face grooming, in the course of early ontogeny.
"The morphogenesis of stereotyped behavior induced by the dopamine receptor agonist apomorphine in the laboratory rat."
H Szechtman, K Ornstein, P Teitelbaum and I Golani (1985)
Neuroscience. Vol. 14 (3) , pp. 783-98.
Abstract: The seemingly unrelated stereotyped locomotor "acts" reported in the literature to be produced by apomorphine in rats are shown to be composites, whose form and sequence are determined by the particular values of a few component variables which form a common denominator in each of the behaviors. Three variables, continuous snout contact, forward progression and turning, account for much of the behavior. In the course of the drug's action these emerge in succession and vary in amount, the latter two successively reaching a peak and subsiding. The interaction between forward progression and turning yields in sequence, forward walking, circling, revolving, tight pivoting and finally side-to-side movements of the forequarters around the relatively stationary hindquarters. Later behaviors in this list are gradually incorporated into the sequence as earlier ones are eliminated. The course of change in forward progression and turning is also reflected in changes in the sequence and in the direction of stepping of each of the four legs. The order in which the behavior unfolds under the drug is opposite to that manifested in ontogeny and in recovery from lateral hypothalamic damage, suggesting that at the particular high dose used, apomorphine is acting not only to activate the behavior but also to shut it down.
"A motility-immotility gradient in the behavior of the inferior wolf during ritualized fighting"
Ilan Golani, Greg Moran and D G Kleiman (1983)
, In Advances in the study of mammalian behavior. Vol. Special Pu (7) , pp. 65 - 94.
"Snout contact fixation, climbing and gnawing during apomorphine stereotypy in rats from two substrains"
H Szechtman, K Ornstein, P Teitelbaum and I Golani (1982)
European journal of pharmacology. Vol. 80 (4) , pp. 385-392.
Abstract: Apomorphine, at doses greater than or equal to 10 mg/kg (intraperitoneally), produced two patterns of stereotypy. In rats from one supplier it induced predominantly gnawing while in those from another predominantly climbing, suggesting that the response to the drug is influenced by genetic and/or experimental factors. At lower doses, apomorphine induced climbing in both groups (ED50 = 1.4 mg/kg in each group) but oral behavior in only one of them (ED50 = 1.3 mg/kg in one, and 8 mg/kg in the second group). Thus, at a given dose of apomorphine, different patterns of stereotypy may result from an interaction between two phenomena: the relative setting of the thresholds to mouth and to climb, and an inverse relation between oral activity and climbing. Analysis of climbing suggests that this response is comprised of two (previously unidentified) fundamental effects of apomorphine: snout contact fixation and bodywise forward progression.
"" Warm-up" along dimensions of movement in the ontogeny of exploration in rats and other infant mammals"
Ilan Golani, G Bronchti, D Moualem and P Teitelbaum (1981)
Proceedings of the National Academy of Sciences of the United States of America. Vol. 78 (11) , pp. 7226-7229.
Abstract: When some infant mammals are placed outside their nest, a sequence of exploratory behavior occurs, displaying a regular buildup and spread of activity. This "warm-up" involves repetition of movement along specific dimensions and an orderly transition from one dimension to the next, with cephalocaudal recruitment of body and limb segments. A similar principle of organization applies to neurological recovery from lateral hypothalamic akinesia.
"A description of relational patterns of movement during ‘ritualized fighting’ in wolves"
Greg Moran, John C. JC John C. Fentress and Ilan Golani (1981)
Animal Behaviour. Vol. 29 (4) , pp. 1146-1165.
Abstract: Schenkel (1947, 1967) was the first to describe ‘ritualized fighting’ in wolves. The current study identifies a set of movement patterns employed during such interactions. The relations between the interactants' movements and the contribution of each individual to these relations are emphasized. Three relational variables are employed to describe interactions: relative distance, relative orientation, and the points of opposition between the interactants. These variables in combination form a three-dimensional interaction space in which a single point describes the momentary state of the configuration of the wolves. The maintenance of four relative configurations plus five transitions between such configurations comprised a consistent set of behavioural patterns. These regular patterns of relational movements indicate that each interactant's movements are constrained in part by a set of rules related to the simultaneous movements of the social partner. In addition, the description of the actual movements in the environment by the individual interactants revealed role-dependent individual contributions by the two interactants to the coordination and management of the relational variables.
"Coordination and integration in the hindleg step cycle of the rat: Kinematic synergies"
Tel Aviv, Ilana Ganor and Ilan Golani (1980)
Brain Research. Vol. 195 (1) , pp. 57-67.
Abstract: The kinematics of the hindleg step cycle of the rat in the vertical domain is composed of 7 synergies. Additionally, a global spatiotemporal principle ensures that each segment of the leg is never displaced backwards. The concepts of flexion and extension are inadequate for the description of step kinematics. As a limb segment changes its orientation, it does so in relation to the next serially connected limb segment or else in relation to gravitation. We call these two aspects of kinematics ‘movements’ and ‘displacements’ respectively, and describe segment kinematics in these terms. Of the 7 kinematic synergies, 5 involve a specific invariant interplay between ‘movements’ and ‘displacements’. Together with the two additional parts they form the skeleton around which the step is organized. The flexible and regulatory nature of the step is obtained by the superposition of biasable properties on top of this skeleton. These include the durations, amplitudes, and initial and final positions of movements-displacements The formalization of the step cycle kinematics represents explicitly intralimb coordination and integration. It also specifies the demand made upon the muscular and neural background organization that mediates the kinematics in a language which is appropriate for neurophysiological investigation.
"A proposed natural geometry of recovery from akinesia in the lateral hypothalamic rat"
I Golani, DL Wolgin and P Teitelbaum (1979)
Brain research. Vol. 164 (1-2) , pp. 237--267.
Abstract: The Eshkol-Wachmann movement notation is used to analyze and describe neurological recovery from the akinesia caused by severe bilateral lateral hypothalamic (LH) damage in rats. Exploratory movement recovers along several relatively independent dimensions which appear successively. First, lateral head scanning movements recover. At about the same time or later, longitudinal (backward-forward) head scans appear. After movements along these two dimensions increase in amplitude and involve the whole body, vertical (dorsal-ventral) head scans with snout contact (along vertical surfaces) typically appear, and increase gradually in amplitude. Later, vertical rearing without snout contact emerges. Recovery proceeds cephalocaudally, as more caudal limb and body segments are recruited along each of the above dimensions separately. LH rats show delayed recruitment of caudal limb and body segments (‘strait-jacket ohenomenon’). Support of the body and management of limb and body segments' contact with the ground also recover relatively independently, in a proximodistal fashion. In recovery, arrests between bouts of activity become shorter. Movement first becomes organized in relation to the animals' own body, and only much later, in relation to the environment. In each sequence of movement after pronounced immobility, the rat recapitulates the process of recovery; and, any time it starts to move, it repeats the movements at a particular amplitude several times until there is an increase to the next larger size movement (‘warm-up’ phenomenon). These regularities explain the apparently bizarre stereotypes behavior in partial enclosures (behavioral traps) seen in LH rats recovering from akinesia. They also explain some aspects of exploration in rats and normal social behavior of wild animals, particularly in situations involving fear and conflict.
"Homeostatic motor processes in mammalian interactions: a choreography of display"
Ilan Golani (1976)
, In Perspectives in ethology. , pp. 69--134.
Abstract: The Eshkol-Wachmann movement notation is used to describe motor sequences in the interactions of both golden jackals (Canis aureus) and Tasmanian devil (Sarcophilus harrisi).
Annotation: From Duplicate 1 ( Homeostatic motor processes in mammalian interactions: a choreography of display - Golani, Ilan )

@incollectiongolani1976homeostatic, title=Homeostatic motor processes in mammalian interactions: a choreography of display,
author=Golani, Ilan,
booktitle=Perspectives in ethology,
pages=69--134,
year=1976,
publisher=Springer

"Non-Metric Analysis of Behavioral Interaction Sequences in Captive Jackals (Canis Aureus L.)"
Ilan Golani (1973)
Behaviour. Vol. 44 (1) , pp. 89-112.
Abstract: A new method is used for the analysis of precopulatory display of jackals. Behavior patterns are described as sequences of configurations. Each configuration is composed of discrete, simultaneous events, such as specific car position, specific tail position, specific body position, etc. With the aid of a non-metric computer technique, it is demonstrated that the recurrence of any particular configuration of behavioral events, performed by a pair of jackals during display, is extremely rare (30 out of 2000). In spite of this heterogeneity of configurations, there is, within short periods of time, some regularity in the composition of the events which form a configuration. This paper explores the question, to what degree the events that form a configuration are discrete to the jackals themselves, as distinguished from the observer, i.e. to what degree they possess a particular significance to the animals, and what is the nature of this significance. The significance of a specific event is defined in this paper as its relatedness to other events both in the configuration and in the temporal sequence. The permanent change in the regularity of composition implies a permanent change in the significance of specific events. This indicates that it is necessary to trace the nature of the change from one significance to another, rather than to look for stable, unchanging significances.
"Sequences of Precopulatory Behavior of the Jackal (Canis Aureus L.)"
I. Golani and H. Mendelssohn (1971)
Behaviour. Vol. 38 (1) , pp. 169-191.
Abstract: The present paper presents a first order description of the precopulatory behavior of the golden jackal (Canis aureus syriacus) without using emotionally-toned words. The coarse structure of this behavior which extends over 4 months prior to copulation is described. During this time, jackals roam the fields in pairs, each pair in a particular area. From time to time they perform certain series of actions while circling each other in a specific manner. The term "T-sequence" is used for this behavior which might be defined as precopulatory. A detailed fine-grain description of selected actual sequences of four captive pairs and three wild pairs is presented. Common features and variation in performance, with special emphasis on the latter, are summarized. Precopulatory behavior of jackals might be viewed as a complex of three groups of events superimposed on one another: the group of events which constitute the T-sequences (such as circling, standing perpendicularly to the mate, putting the head on the mate's back, etc.) is superimposed on a group of patterns which are performed during the first phase of precopulatory behavior - sniffing the ground, urination and scraping. Later, a third group of patterns is superimposed on the first two groups: this group includes the male licking the female's vulva, the female mounting the male and other patterns. At this stage, the frequency of appearance of patterns of the first two groups decreases gradually; the male starts to mount the female, and a few days later copulations follow. Variation in performance was found in the following acts: 1) In certain pairs the male always forms the top of the T during T-sequences, in others the female, and in still other pairs both mates take turns at forming either the top or the base of the T. Relative body position during T-sequences is in correlation with the distribution of events between the mates. It also affects the time of onset of the week of copulations. 2) Some "agonistic" patterns were observed during T-sequences only in captivity. 3) Certain behavior patterns are performed by some pairs but not by others. 4) Some behavior patterns are performed in different "styles" by different individuals. The performance of T-sequences is related to the following factors: a) sequential relation of events to copulation, b) time of day, c) change in environment, d) howling, e) duration of pair bond, f) individual differences. Motivational models as well as quantitative analysis were avoided, since the variability of the material calls for the development of special methods for fine-grain analysis. This material deserves phenomenological treatment, in the sense that regularity should be looked for first in the structure of concrete specific behavior sequences, and only then in the behavior in general.