Behavior Research Methods

, Volume 50, Issue 4, pp 1673–1685 | Cite as

Methods for the effective study of collective behavior in a radial arm maze

  • Johann Delcourt
  • Noam Y. Miller
  • Iain D. Couzin
  • Simon Garnier


Collective behaviors are observed throughout nature, from bacterial colonies to human societies. Important theoretical breakthroughs have recently been made in understanding why animals produce group behaviors and how they coordinate their activities, build collective structures, and make decisions. However, standardized experimental methods to test these findings have been lacking. Notably, easily and unambiguously determining the membership of a group and the responses of an individual within that group is still a challenge. The radial arm maze is presented here as a new standardized method to investigate collective exploration and decision-making in animal groups. This paradigm gives individuals within animal groups the opportunity to make choices among a set of discrete alternatives, and these choices can easily be tracked over long periods of time. We demonstrate the usefulness of this paradigm by performing a set of refuge-site selection experiments with groups of fish. Using an open-source, robust custom image-processing algorithm, we automatically counted the number of animals in each arm of the maze to identify the majority choice. We also propose a new index to quantify the degree of group cohesion in this context. The radial arm maze paradigm provides an easy way to categorize and quantify the choices made by animals. It makes it possible to readily apply the traditional uses of the radial arm maze with single animals to the study of animal groups. Moreover, it opens up the possibility of studying questions specifically related to collective behaviors.


Collective behavior Collective decision-making Group cohesion Fission–fusion societies 


Author note

J.D. is a postdoctoral researcher at the Fonds de la Recherche Scientifique (FRS)–FNRS (Belgium). This work was supported by the FRS–FNRS under FRFC Grants 2.4617.08F, 2.4507.08.F, and T.1064.14 (PDR FNRS project). I.D.C. acknowledges support from the NSF (Grants PHY-0848755, IOS-1355061, and EAGER-IOS-1251585), ONR (Grants N00014-09-1-1074, N00014-14-1-0635), ARO (Grants W911NG-11-1-0385, W911NF-14-1-0431), and the Human Frontier Science Program (RGP0065/2012). We thank Adrian de Froment for useful discussions and for helping with part of the experiments. We also thank Pascal Poncin and Jean-Louis Deneubourg for their support and advice in the PDR FNRS Project. We thank C. Orban for her advice on the writing.

Supplementary material

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ESM 1 (DOCX 70 kb)
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Fig. S1. Illustration of transitions of majority. (a) Transition of the majority between arm 1 and arm 3. A majority is observed until time t1 in arm 1, a new majority group is observed in arm 3 from time t3. Between the times t1 and t3, no majority is observed in any arm (the central zone is not considered). The period of transition between two successive majority periods (and its duration) is defined as being between t1 and t3. (b) Theoretical case where, after a majority period (here in arm 1 in time t1), individuals move into different arms without achieving a new majority state (at time t2 in this example). A new majority is achieved at time t3 after reunification of two small groups coming from arms 3 and 4. Note, in this example, that the new majority does not involve all the individuals of the previous majority, one individual remaining in arm 5. “M” indicates the “majority arm”. See Fig.1 for definition of arm number and majority arm. (DOCX 230 kb)
13428_2018_1024_MOESM4_ESM.docx (99 kb)
Fig. S2. Theoretical example illustrating the concept of different types of majority transition. Upper panel: a timeline with periods of majority, M (indicated by color blocks; each color corresponds to a specific arm), alternating with periods of no majority, NM (uncolored blocks). The state sequence of observed majority is the sequence of arm identities (indicated by upper-case letters) where a majority is successively observed. The duration of the majority state is not taken into account. This sequence constitutes the first order transitions of majority. Second order transitions of majority indicate the sequence of transitions between a majority and the second-next majority achieved (in time). Majority transition without repetition is defined to filter cases where transition has aborted, and is based on only the first order transition: the first order transitions between two successive identical arms are ignored. For higher order transition, repetitive identical majority are considered as only one element in the state sequence. (DOCX 99 kb)
13428_2018_1024_MOESM5_ESM.docx (234 kb)
Fig. S3. Several theoretical examples of stereotypic motion patterns (algorithmic behaviors) with corresponding majority transition diagrams for first and second order transitions. The numbers are the expected percentage of choice of the majority for each majority transition (and underlined also by the thickness of the transition arrows). (a) case where the majority of the group always chooses the next arm to the left; (b) case where the majority oscillates between only two arms; (c) case of a “star” pattern where the majority moves from one particular arm to any other arm, and returns immediately afterwards to the initial majority arm; (d) case in which the majority moves only into the adjacent left or right arm. (DOCX 234 kb)
13428_2018_1024_MOESM6_ESM.docx (263 kb)
Fig. S4. Distribution of the number of scorings performed by human counters on each of the 180 test images. The dotted line is the median of the distribution. (DOCX 263 kb)
13428_2018_1024_MOESM7_ESM.docx (249 kb)
Fig. S5. Example of a sequence of 30 minutes, illustrating the relationship between maximal group size observed in the maze arms (the central zone is not included) and periods with and without majority. The group size was ten fish. Each vertical colored bar shows a period during which there was a majority in a specific arm; the color indicates which arm. Between each period of majority is a period of transition during which there was no majority in any arm (n < 6; uncolored gaps). The dotted line is the minimum group size required to obtain a majority in an arm (n ≥ 6), independent of the central zone. (DOCX 248 kb)


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Copyright information

© Psychonomic Society, Inc. 2018

Authors and Affiliations

  • Johann Delcourt
    • 1
  • Noam Y. Miller
    • 2
  • Iain D. Couzin
    • 3
    • 4
  • Simon Garnier
    • 5
  1. 1.Behavioural Biology UnitUniversity of LiègeLiègeBelgium
  2. 2.Department of PsychologyWilfrid Laurier UniversityWaterlooCanada
  3. 3.Department of Ecology and Evolutionary BiologyPrinceton UniversityPrincetonUSA
  4. 4.Department of Collective Behaviour, Max Planck Institute for Ornithology and Department of BiologyUniversity of KonstanzKonstanzGermany
  5. 5.Department of Biological SciencesNew Jersey Institute of TechnologyPrincetonUSA

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