Nature Vol. 254 Aprit 10 1975 Aggression and mating success in Drosophila melanogaster ‘Tue value of Drosophila. melanogaster as_an experimental animal for the study of the genetics of behaviour increases as more specific behaviours are shown to exist this species. Although aggresive behaviour has been reported. for_D. Subobscura! and some Hlawalian® species, and has been briefly mentioned for many other species, it has not yet been des- Gribed for D. melanogaster. We have repeatedly observed fochaviours which we. interpreted as aggressive and here, report an experiment designed to investigate aggression and ite relationship to mating success—an important component of fines. Flies from a mass-bred stock (Haren) which has been in the laboratory since July, 1971 were used in this experiment. To identify individuals, 12-vold males were marked on’ the oral surface of the thorax with dots of acrylic paint while tinder light etherisation, They were Kept singly in vials con- taining standard. Drosophila mediam until they were 3 4 ol. ‘AY ths time, six males were placed in a eslindrical Perspex arena (7 em diameter, 4 cm high) containing a small dish G5 cm diameter) with Drosophila medium. After an 18h seiling perio, ies were observed with a binocular microscope. ‘All interactions occurring on the fuod surface and their out- comes were recorded over & period of 3-8. In addition the Fig. 1A charging male of D. melanogaster: wing threst, wings spread, raised and (wisted. 1 ait asiz Fig. 2 Two males boxing. position of the individuals in relation to the food surface was ‘was noted every 5 min, We then started to introduce 3-d-old virgin females at intervals of approximately $0 min, recording, which males were successful in copulation. At the end of each day the females were removed. ‘The interactions observed were primarily aggressive or sexual. We found three frequently occurring aggressive be- haviour patterns: wing threat (Fig. 1) which is often directed to ‘other males just before avery quick charge, in which the aggressor usually rises on his hindleys shortly before impact, is made, and boxing (Fig. 2) which comprises several variations of very’ vigorous slashing and tapping with the front legs, often while both males rise on their hindlegs. ae i} | fq bale fh nono le oe Fig. 3 Activities of each lo expressed ss percentages. (1) Time spent on the food, 100%," tolal time of observation: (2) attacks initated, 100% otal number of attacks made by all fies: G)'Rghts won, 106% total of a given animal's tights: () coputations, 1007," total number of copulations by all males= 19 ‘The six males showed large differences in the frequency with “which they stayed on the food surface, with which they attacked, ‘were attacked, won and lost bouts of aggression and copulated (Fig. 3). The’ winner is defined as the animal which remains fon the food surface after an aggressive encounter. Clearly, ‘white’ stayed on the food surface most often (x? 64.1 5 df, P<0.005), initiated most threats and attacks relative to the amount of time spent on the food (*=31.0, 3 , P-<0,005) and won 100% of his fights. Ranking the animals o' the proportion of fights won gives the following result: white’ ~ ‘green’ > “blue I' > ‘red’ > “blue 2'~'yellow’. This rank order is significantly correlated (r=0.95, P<0.01) with the ‘number of copulations achieved by each male Drosophila males do court other males, and other invest gators who have encountered the behaviours we have des- cribed may have interpreted them as sexual in nature. In well over 1,000 videotaped single pair courtships, however, none Of the’ behaviour patterns that we define as aggressive has previously been observed. Milani* has described @ behaviour Nature Vol. 254 April 10 1975 similar to our wing threat in D. subobscura and interpreted it as fggressive in nature, Brown’ dismisses this notion and prefers to call it counter-signalling. According to our observations, however, it cannot be a counter signal since a male engages in this behaviour before an interaction initiated by him. In D. melanogaster, wing flicking serves as a counter signal which is given in interactions involving physical contact between individuals of either sex. ‘The adaptive nature of aggression on a food source might be that males monopolise a food resource for reasons of energy requirement and/or because females will be attracted to it ‘Our resulls provide evidence that the more successful a male is in fights the fitter he is in terms of mating success, ‘This work was supported by scholarships from the National Research Council of Canada and the German Academie EXx- ‘change Service. We thank Professor A. Manning and Mr R. ‘Weight for their helpful criticisms of the manuscript and Mr D. Cremer for his assistance with the photography. Maurice A, Dow FLORIAN VON SCHILCHER Department of Zoology, University of Edinburgh, West Mains Road, Edinburgh EH9 307, UK Restved December 3, 19945 reve February 14, 1975, " Mig R. Seed Sonic Paes, ao Sepere Sait, 1, 21%-28 2 splethe HL, Up Tex, Publ 6, 285-313 (1966) 2 Shh ee Bal de ae Nel ae 9p 8s" 474 98 4 Browne G2 hair 236-1081 0960),