Laboratory of Animal Behavior

Ilan Golani,
Professor of Zoology,
Department of Zoology,
Faculty of Life Sciences,
Tel Aviv University,
Tel Aviv,

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Research in the interface between

Behaviour, Computation, Statistics and Genetics

We presently focus on two projects:

  1. Motor coordination. Our current treatment of this subject is illustrated in two recent papers that can be downloaded below (Kafkafi et al., 1996; Kafkafi and Golani, 1998), and by two animations, prepared by Neri Kafkafi, which accompany the second of the two papers ("A traveling wave of lateral movement coordinates...."). The first animation explains the application of the concept of relative phase to unrestrained locomotor behavior. You would first have to download  Mathreader and then click Animation of relative phase(3.6 Mb).
    The second animation shows that ferret locomotor behavior involves a travelling, rather than a standing wave of lateral movement. This is accomplished by coloring each of the parts of the body of the animated animal in correspondence with its current phase of movement. To see it click   Phase animation in ferret. (2.2 Mb).

  2. Algorithmic definitions of behavior patterns in mice and their utility in genetic research. Collaborative research is  performed in Tel Aviv University (TAU) and the Maryland Psychiatric Research Center (MPRC) in Baltimore.  In TAU it is performed by a team including  zoologists and statisticians. The project involves the use of large data bases and scientific visualization. For further information write to:  see also Prof. Yoav Benjamini, and Dr. Neri Kafkafi .

For details on current projects see:

  1. SEE: a tool for the visualization and analysis of rodent exploratory behavior.
  2. Phenotyping mouse exploratory behavior (2.2 Mb).

Our research on mouse behavior:

The focus in the next stages of genomic research will undoubtedly be on comparative analysis of genomes. "Expression" and "function" are the watch words of the new era. Perhaps the most intriguing question in this quest is that of the relationship between genomes and behavioral phenotypes. A major stumbling block in this quest, however, is the lack of analytic tools on the behavior side of the genome/behavior interface. Ethology ,neuroethology, behavioral pharmacology and behavioral neuroscience failed to develop a methodology that can handle the representation of free whole-animal behavior at a level of articulation and rigor be fitting that employed on the genome side.

We are a research team currently engaged in developing such a methodology. Our team  includes several zoologists (Ilan Golani, Dina lipkind, Ehud Fonio, Guy Horev, Eyal Gruntman, Anna Dvorkin, Eran Polosetski, Tel Aviv, and Neri Kafkafi, Baltimore, The Maryland Psychiatric Research Institute,  two statisticians (Yoav Benjamini, and Anat Sakov, Tel Aviv),  and a neurobiologist ( Greg Elmer, The Maryland Psychiatric Research Institute, Baltimore). Because the mouse has now become the experimental animal par excellence of Behavior Genetics, our work is centered on mice. In work already done by us and by others, it has been shown that contrary to what was common belief, rodent free locomotor behavior is quite structured. In the last few years we have developed interactive software for analyzing this structure. This software can visualize and quantify the patterns, using as input the automatically digitized time-series of the animal's location (for a video clip illustrating the tracking process of the behavior of a rat in a large 6.5m diameter arena see "Tracking of rat exploratory behavior" (28.5 Mb) and producing as output, key parameters that characterize the behavior. These parameters are relatively independent of each other, and reveal a natural structure that is relatively independent of the animal's level of activity. They reflect processes involving motivation, navigation, spatial memory and learning. They can be measured automatically and efficiently in very large numbers of mice (high throughput) and their listing should result in an objective algorithmic definition of species- and strain-specific behavior.

Having at hand a long list of numerical values that summarize structural differences between genetically distinct mouse strains whose genetic map is at a relatively advanced stage of construction introduces two statistical problems: that of the False Discovery Rate (FDR; Benjamini and Hochberg, 1995) stemming from the simultaneous performance of multiple comparisons (by increasing the probability of apparently significant differences), and that of using the Quantitative Trait Loci (QTL) approach, in order to map the location of genes having specific behavioral function on the chromosome. Studying these multiple relations between a large number of  locations on the genome and long lists of behavioral endpoints require an extensive interaction between the members of our group.

Our laboratory is fully equipped to perform state of the-art behavior research, and  works closely with the Maryland Psychiatric Institute and the Tennessee Mouse Genome Consortium, to produce a diverse and vibrant research environment. Our mission includes the creation of large databases available to all, and the development of analytical tools to support their use.


Controlling the false discovery rate in behavior genetics research.
Behavioural Brain Research Volume 125,Issues 1-2
8 November 2001, Pages 279-284
Yoav Benjamini, Dan Drai, Greg Elmer, Neri Kafkafi and Ilan Golani
The journal

Rats and mice share common ethologically relevant parameters of exploratory behavior.
Behavioural Brain Research Volume 125, Issues 1-2
8 November 2001 Pages 133-140
Dan Drai, Neri Kafkafi, Yoav Benjamini, Greg Elmer and Ilan Golani
The journal

SEE: a tool for the visualization and analysis of rodent exploratory behavior.
Neuroscience & Biobehavioral Reviews, 25 (2001) 5,
Pages 409-426
Dan Drai and Ilan Golani
The journal

Natural segmentation of the locomotor behavior of drug-induced rats in a photobeam cage.
Journal of Neuroscience Methods 109 (2): 111-121 AUG 30 2001
Neri Kafkafi, Cheryl Mayo, Dan Drai, Ilan Golani and Greg Elmer

Phenotyping stereotypic behaviour: collective variables, range of variation and predictability.
Volume 65 (1999) 191-220 Applied Animal Behavior Science
I. Golani, N.Kafkafi, D. Drai.

The dynamics of long-term exploration in the rat
Biological Cybernetics, Jul 20, 1998 Vo78, Issue 6, pp 423-432
Ofer Tchernichovski , Yoav Benjamini , Ilan Golani

The dynamics of long term exploration in the rat
Ofer Tchernichovski , Yoav Benjamini
Volume 78, Issue 6, pp 433-440
Biological Cybernetics, Jul 20, 1998
Download article in PDF format here

Statistical discrimination of natural modes of motion in rat exploratory behavior
Dan Drai, Yoav Benjamini, Ilan Golani
Volume (issue): 96 (2) 2000
Journal of Neuroscience Methods
Download article in PDF format here

(For a video clip illustrating the phenomenon which is the subject of this paper: alternation between progression and stayingin place behavior click "alternation between lingering and progression in exploratory behavior of a wild Rattus norvegicus "(27.5 Mb)).

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 , Yoav Benjamini
Volume 74, Issue 6, pp 487-495
Biological Cybernetics, Feb 14, 1996
Download abstract here

A traveling wave of lateral movement coordinates both turning and forward walking in the ferret
Neri Kafkafi , Ilan Golani
Volume 78, Issue 6, pp 441-453
Biological Cybernetics, Jul 20, 1998
Download article in PDF format here

Controlling the False Discovery Rate: a practical and powerful approach to multiple testing.
Benjamini Y, Hochberg Y (1995)

Journal of the Royal Statistical Society, SeriesB, 57, 1, 289-300.


A mobility gradient in the organization of vertebrate movement: The perception of movement through symbolic language.
Golani I (1992) 

SEE locomotor behavior test discriminates C57BL/6J and DBA/2J mouse inbred strains across laboratories and protocol conditions
Kafkafi N, Lipkind D, Benjamini Y, Mayo CL, Elmer GI, Golani I



Darting behavior: a quantitative movement pattern designed for discrimination and replicability in mouse locomotor behavior
Kafkafi N, Pagis M, Lipkind D, Mayo CL, Benjamini Y, Golani I, Elmer GI

BEHAVIOURAL BRAIN RESEARCH 142 (1-2): 193-205 JUN 16 2003


The Dynamics of Spatial Behavior: How can robust smoothing techniques help?
Hen I, Sakov A, Kafkafi N, Golani I, Benjamini Y

JOURNAL OF NEUROSCIENCE METHODS 133 (1-2): 161-172 FEB 15 2004


New replicable anxiety-related measures of wall vs. center behavior of mice in the open field
Lipkind D, Sakov A, Kafkafi N, Elmer GI, Benjamini Y, Golani I



Locomotor and exploratory behaviour. In The Behavior of the Laboratory Rat A Handbook with Tests Edited by Ian Q. Whishaw and Bryan Kolb.
Golani I, Benjamini Y, Dvorkin A, Lipkind D, Kafkafi N

Oxford University Press.


Genotype-Environment Interactions in Mouse Behavior: A Way Out of the Problem
Kafkafi N, Benjamini Y, Sakov A, Elmer G, Golani I Proceedings of the National Academy of Sciences of the United States of America 102.12 (2005): 4619-4624.?


New replicable anxiety-related measures of wall vs. center behavior of mice in the open field

Lipkind D, Sakov A, Kafkafi N, Elmer GI, Benjamini Y, Golani I. JOURNAL OF APPLIED PHYSIOLOGY 97 (1): 347-359 JUL 2004

Activity density in the open field: a measure for differentiating the effect of psychostimulants.


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, Ilan Golani. PLoS ONE 01/2013; 8(10):e76257

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.

Measuring behavior of animal models: faults and remedies.

Ehud Fonio , Ilan Golani, Yoav Benjamini. Nature Methods 12/2012; 9(12):1167-1170.

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. Behavioural brain research 01/2012; 231(2):309-16.

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.

Short and long term measures of anxiety exhibit opposite results.

Ehud Fonio , Yoav Benjamini, Ilan Golani. PLoS ONE 01/2012; 7(10):e48414.

This is a twin article, to be read before the "Nature Methods" article " Measuring behavior of animal models: faults and remedies".

Validation of the dimensionality emergence assay for the measurement of innate anxiety in laboratory mice.

Apar Jain , Anna Dvorkin,Ehud Fonio, Ilan Golani, Cornelius T Gross. European neuropsychopharmacology: the journal of the European College of Neuropsychopharmacology 07/2011; 22(2):153-63. ·

Quantifying the buildup in extent and complexity of free exploration in mice.

Yoav Benjamini , Ehud Fonio,Tal Galili, Gregor Z Havkin, Ilan Golani

Proceedings of the National Academy of Sciences 03/2011; 108 Suppl 3:15580-7.

High-throughput data analysis in behavior genetics

Anat Sakov , Ilan Golani, Dina Lipkind,Yoav Benjamini. The Annals of Applied Statistics 11/2010; ·

Ten ways to improve the quality of descriptions of whole-animal movement.

Yoav Benjamini , Dina Lipkind,Guy Horev,Ehud Fonio, Neri Kafkafi, Ilan Golani

Neuroscience & Biobehavioral Reviews 07/2010; 34(8):1351-65.

Superiority and Inferiority: a Morphological Analysis of Free and Stimulus Bound Behaviour in Honey Badger (Mellivora capensis) Interactions

Yona Yaniv, Ilan Golani. Ethology 04/2010; 74(2):89 - 116. ·

Knots: attractive places with high path tortuosity in mouse open field exploration.

Anna Dvorkin , Henry Szechtman, Ilan Golani PLoS Computational Biology 01/2010; 6(1):e1000638.

Freedom of movement and the stability of its unfolding in free exploration of mice.

Ehud Fonio , Yoav Benjamini, Ilan Golani. Proceedings of the National Academy of Sciences 11/2009; 106(50):21335-40.

Mouse cognition-related behavior in the open-field: emergence of places of attraction.

Anna Dvorkin , Yoav Benjamini, Ilan Golani. PLoS Computational Biology 03/2008; 4(2):e1000027.

Characterizing Animal Behavior through Audio and Video Signal Processing

D. Valente , Haibin Wang,P. Andrews,P.P. Mitra,S. Saar,O. Tchernichovski, I. Golani, Y. Benjamini. IEEE Multimedia 11/2007; ·

Coordination of steering in a free-trotting quadruped.

Eyal Gruntman , Yoav Benjamini, Ilan Golani. Journal of Comparative Physiology 04/2007; 193(3):331-45.

Estimating wall guidance and attraction in mouse free locomotor behavior.

G Horev , Y Benjamini, A Sakov, I Golani

Genes Brain and Behavior 03/2007; 6(1):30-41.

Analysis of the trajectory of Drosophila melanogaster in a circular open field arena.

Dan Valente , Ilan Golani, Partha P Mitra

PLoS ONE 02/2007; 2(10):e1083.

Characterization of animal behavior through the use of audio and video signal processing .

Dan Valente , Haibin Wang,Peter Andrews,Sigal Saar,Ofer Tchernichovski, Ilan Golani,Yoav Benjamini, Partha Mitra. IEEE Multimedia - IEEEMM. 01/2007; 14(99).

Wild mouse open field behavior is embedded within the multidimensional data space spanned by laboratory inbred strains. E Fonio , Y Benjamini,A Sakov, I Golani. Genes Brain and Behavior 08/2006; 5(5):380-8.

Multimedia signal processing for behavioral quantification in neuroscience.

Peter Andrews , Haibin Wang,Dan Valente,Jihène Serkhane,Partha P. Mitra,Sigal Saar, Ofer Tchernichovski, Ilan Golani. Proceedings of the 14th ACM International Conference on Multimedia, Santa Barbara, CA, USA, October 23-27, 2006; 01/2006

Genotype-environment interactions in mouse behavior: a way out of the problem.

Neri Kafkafi , Yoav Benjamini,Anat Sakov, Greg I Elmer, Ilan Golani