Modeling shortest path selection of the ant Linepithema humile using psychophysical theory and realistic parameter values.

The emergence of self-organizing behavior in ants has been modeled in various theoretical approaches in the past decades. One model explains experimental observations in which Argentine ants (Linepithema humile) selected the shorter of two alternative paths from their nest to a food source (shortest path experiments). This model serves as an important example for the emergence of collective behavior and self-organization in biological systems. In addition, it inspired the development of computer algorithms for optimization problems called ant colony optimization (ACO). In the model, a choice function describing how ants react to different pheromone concentrations is fundamental. However, the parameters of the choice function were not deduced experimentally but freely adapted so that the model fitted the observations of the shortest path experiments. Thus, important knowledge was lacking about crucial model assumptions. A recent study on the Argentine ant provided this information by measuring the response of the ants to varying pheromone concentrations. In said study, the above mentioned choice function was fitted to the experimental data and its parameters were deduced. In addition, a psychometric function was fitted to the data and its parameters deduced. Based on these findings, it is possible to test the shortest path model by applying realistic parameter values. Here we present the results of such tests using Monte Carlo simulations of shortest path experiments with Argentine ants. We compare the choice function and the psychometric function, both with parameter values deduced from the above-mentioned experiments. Our results show that by applying the psychometric function, the shortest path experiments can be explained satisfactorily by the model. The study represents the first example of how psychophysical theory can be used to understand and model collective foraging behavior of ants based on trail pheromones. These findings may be important for other models of pheromone guided ant behavior and might inspire improved ACO algorithms.

[1]  K. Akre,et al.  Psychophysics and the evolution of behavior. , 2014, Trends in ecology & evolution.

[2]  Lars Chittka,et al.  Speed-accuracy tradeoffs in animal decision making. , 2009, Trends in ecology & evolution.

[3]  Thomas Stützle,et al.  Ant Colony Optimization , 2009, EMO.

[4]  S. Pratt,et al.  Ant colonies outperform individuals when a sensory discrimination task is difficult but not when it is easy , 2013, Proceedings of the National Academy of Sciences.

[5]  E. B. Wilson Probable Inference, the Law of Succession, and Statistical Inference , 1927 .

[6]  J. Deneubourg,et al.  Modulation of trail laying in the antLasius niger (Hymenoptera: Formicidae) and its role in the collective selection of a food source , 1993, Journal of Insect Behavior.

[7]  Anna Dornhaus,et al.  The Trail Less Traveled: Individual Decision-Making and Its Effect on Group Behavior , 2012, PloS one.

[8]  S. K. Park,et al.  Random number generators: good ones are hard to find , 1988, CACM.

[9]  J. Deneubourg,et al.  Trail laying behaviour during food recruitment in the antLasius niger (L.) , 1992, Insectes Sociaux.

[10]  J. Deneubourg,et al.  The self-organizing exploratory pattern of the argentine ant , 1990, Journal of Insect Behavior.

[11]  J. Movshon,et al.  The analysis of visual motion: a comparison of neuronal and psychophysical performance , 1992, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[12]  J. Deneubourg,et al.  Memory and chemical communication in the orientation of two mass-recruiting ant species , 1993, Insectes Sociaux.

[13]  A. Dussutour,et al.  Noise improves collective decision-making by ants in dynamic environments , 2009, Proceedings of the Royal Society B: Biological Sciences.

[14]  F A Wichmann,et al.  Ning for Helpful Comments and Suggestions. This Paper Benefited Con- Siderably from Conscientious Peer Review, and We Thank Our Reviewers the Psychometric Function: I. Fitting, Sampling, and Goodness of Fit , 2001 .

[15]  F. Ratnieks,et al.  Synergy between social and private information increases foraging efficiency in ants , 2011, Biology Letters.

[16]  R Core Team,et al.  R: A language and environment for statistical computing. , 2014 .

[17]  P. L. Robertson,et al.  Exocrine gland involvement in trailing behaviour in the Argentine ant (Formicidae: Dolichoderinae) , 1980, Animal Behaviour.

[18]  J. Spaethe,et al.  Comparative psychophysics of bumblebee and honeybee colour discrimination and object detection , 2008, Journal of Comparative Physiology A.

[19]  Aluizio F. R. Araújo,et al.  Modelling foraging ants in a dynamic and confined environment , 2011, Biosyst..

[20]  W. Hangartner Orientierung vonLasius fuliginosus Latr. an einer Gabelung der Geruchsspur , 1969, Insectes Sociaux.

[21]  Marco Dorigo,et al.  Ant system: optimization by a colony of cooperating agents , 1996, IEEE Trans. Syst. Man Cybern. Part B.

[22]  J. Deneubourg,et al.  Trail-laying behaviour during exploratory recruitment in the Argentine ant: Iridomyrmex humilis (Mayr) , 1989 .

[23]  J. Deneubourg,et al.  How do ants assess food volume? , 2000, Animal Behaviour.

[24]  Dong-Hwan Choe,et al.  Pheromone communication in ants: a detailed analysis of concentration-dependent decisions in three species , 2014, Behavioral Ecology and Sociobiology.

[25]  G. Fechner Elemente der Psychophysik , 1998 .

[26]  S. Key,et al.  Effects of gaster extract trail concentration on the trail following behaviour of the Argentine ant, Iridomyrmex humilis (Mayr) , 1981 .

[27]  William H. Press,et al.  The Art of Scientific Computing Second Edition , 1998 .

[28]  Ernst Heinrich Weber,et al.  De pulsu, resorptione, auditu et tactu. Annotationes anatomicae et physiologicae , 1834 .

[29]  M. Beekman,et al.  A mathematical model of foraging in a dynamic environment by trail-laying Argentine ants. , 2012, Journal of theoretical biology.

[30]  N. Tsutsui,et al.  Trail Pheromone of the Argentine Ant, Linepithema humile (Mayr) (Hymenoptera: Formicidae) , 2012, PloS one.

[31]  Jacobus C. Biesmeijer,et al.  Self-organization in collective honeybee foraging: emergence of symmetry breaking, cross inhibition and equal harvest-rate distribution , 2002, Behavioral Ecology and Sociobiology.

[32]  D. Sumpter,et al.  The role of multiple pheromones in food recruitment by ants , 2009, Journal of Experimental Biology.

[33]  Guy Theraulaz,et al.  Self-Organization in Biological Systems , 2001, Princeton studies in complexity.

[34]  F. Ratnieks,et al.  Combined use of pheromone trails and visual landmarks by the common garden ant Lasius niger , 2008, Behavioral Ecology and Sociobiology.

[35]  E. Wilson Chemical communication among workers of the fire ant Solenopsis saevissima (Fr. Smith) 1. The Organization of Mass-Foraging , 1962 .

[36]  N. Davies,et al.  Characterization of aggregation factors and associated compounds from the argentine ant,Iridomyrmex humilis , 1980, Journal of Chemical Ecology.

[37]  S. Klein,et al.  Measuring, estimating, and understanding the psychometric function: A commentary , 2001, Perception & psychophysics.

[38]  S. Key,et al.  Trail-following responses of the Argentine ant,Iridomyrmex humilis (Mayr), to a synthetic trail pheromone component and analogs , 2004, Journal of Chemical Ecology.

[39]  F. Ratnieks,et al.  Decision making in ant foragers (Lasius niger) facing conflicting private and social information , 2011, Behavioral Ecology and Sociobiology.

[40]  Duncan E. Jackson,et al.  Modulation of pheromone trail strength with food quality in Pharaoh's ant, Monomorium pharaonis , 2007, Animal Behaviour.

[41]  David J. T. Sumpter,et al.  Individual Rules for Trail Pattern Formation in Argentine Ants (Linepithema humile) , 2012, PLoS Comput. Biol..

[42]  J. Deneubourg,et al.  Self-organized shortcuts in the Argentine ant , 1989, Naturwissenschaften.

[43]  Guy Theraulaz,et al.  Path efficiency of ant foraging trails in an artificial network. , 2006, Journal of theoretical biology.