Queen mandibular pheromone: questions that remain to be resolved

The discovery of ‘queen substance’, and the subsequent identification and synthesis of keycomponents of queen mandibular pheromone, has been of significant importance to beekeepers and to thebeekeeping industry. Fifty years on, there is greater appreciation of the importance and complexity of queenpheromones, but many mysteries remain about the mechanisms through which pheromones operate. Thediscovery of sex pheromone communication in moths occurred within the same time period, but in this case,intense pressure to find better means of pest management resulted in a remarkable focusing of research activityon understanding pheromone detection mechanisms and the central processing of pheromone signals in themoth. We can benefit from this work and here, studies on moths are used to highlight some of the gaps in ourknowledge of pheromone communication in bees. A better understanding of pheromone communication inhoney bees promises improved strategies for the successful management of these extraordinary animals

Queen-signal modulation of worker pheromonal composition in honeybees

Worker sterility in honeybees is neither absolute nor irreversible. Whether under queen or worker control, it is likely to be mediated by pheromones. Queen-specific pheromones are not exclusive to queens; workers with activated ovaries also produce them. The association between ovarian activation and queen-like pheromone occurrence suggests the latter as providing a reliable signal of reproductive ability. In this study we investigated the effect of queen pheromones on ovary development and occurrence of queen-like esters in workers' Dufour's gland. Workers separated from the queenright compartment by a double mesh behaved like queenless workers, activating their ovaries and expressing a queen-like Dufour's gland secretion, confirming that the pheromones regulating both systems are non-volatile. Workers with developed ovaries produced significantly more secretion than sterile workers, which we attribute primarily to increased ester production. Workers separated from the queenright compartment by a single mesh displayed a delayed ovarian development, which we attribute to interrupted transfer of the non-volatile pheromone between compartments. We suggest that worker expression of queen-like characters reflects a queen-worker arms race; and that Dufour's gland secretion may provide a reliable signal for ovarian activation. The associative nature between ovary development and Dufour's gland ester production remains elusive. © 2004 The Royal Society.Peer Reviewe

In nest environment modulates nestmate recognition in the ant Camponotus fellah

Multiple behavioral and chemical studies indicate that ant nestmate recognition cues are low-volatile substances, in particular hydrocarbons (HCs) located on the cuticular surface. We tested the hypothesis that in the ant Camponotus fellah, nest environment, in particular nest volatile odors, can modulate nestmate-recognition-mediated aggression. Workers were individually confined within their own nest in small cages having either a single mesh (SM = limited physical contact permitted) or a double mesh (DM = exposed to nest volatiles only) screen. Individual workers completely isolated outside their nest (CI) served as control. When reintroduced into a group of 50 nestmates, the CI workers were attacked as alien ants after only 2 weeks of separation, whereas the SM workers were treated as nestmates even after 2 months of separation. Aggression towards DM ants depended on the period of isolation. Only DM workers isolated for over 2 months were aggressed by their nestmates, which did not significantly differ from the CI nestmates. Cuticular HC analyses revealed that the profile of the non-isolated ants (NI) was clearly distinct from that of CI, SM and DM ants. Profile differences matched the aggressive response in the case of CI ants but were uncorrelated in the case of SM or DM ants. This suggests that keeping the ants within the nest environment affected nestmate recognition in additional ways than merely altering their HC profile. Nest environment thus appears to affect label-template mismatch by modulating aggressive behavior, as well as the direction at which cuticular HCs diverged during the separation period. © Springer-Verlag 2004.Peer Reviewe

Odour convergence and tolerance between nestmates through trophallaxis and grooming in the ant Camponotus fellah (Dalla Torre)

Social isolation provides a useful tool to study nestmate recognition in ants. In Camponotus fellah, reintroduction of 10-day isolated (IS) workers to their colony resulted in intensive trophallaxis and grooming, while longer isolation periods generally provoked rejection of the IS ants. In the first experiment the behaviour of queenless (QL) and queenright (QR) workers towards 10-day IS workers was tested. Trophallaxis of QL or QR with IS workers was of similar magnitude, but was significantly higher than that among the QL or QR, or that between QL and QR workers. Allogrooming was mostly initiated by the resident non-isolated ants (QL or QR) possibly because they detected a slight mismatch between the IS ant's odour and their own template, which represents the group odour. It appears that the presence/absence of the queen did not affect nestmate recognition cues of workers. The second experiment demonstrated that 20-day IS workers were strongly aggressed by colony guards, irrespective of whether they were QL or QR. However, if they were permitted to exchange trophallaxis and grooming with 5 young nestmates (companion ants) for 5 days before reintroduction to their colony, aggression was greatly reduced, irrespective of the origin of the companion ants (QR or QL). Chemical analysis showed a significant divergence between the hydrocarbon profiles of IS and both QL and QR groups, but a prior contact of the IS workers with companion ants resulted in re-convergence of their profile with that of the colony. These results demonstrate that nestmate recognition cues are exchanged between workers via trophallaxis and grooming and that they are not dominated by queen cues, two conditions that fulfil Gestalt nestmate recognition signals requirements.Peer Reviewe

Nest volatiles as modulators of nestmate recognition in the ant Camponotus fellah

When ants from alien colonies encounter each other the stereotypic reaction is usually one of aggressive behavior. It has been shown that factors such as queen-derived cues or nest-odors modulate this reaction. Here we examined whether nest volatiles affect nestmate recognition by observing the reaction of nestmates towards individual workers under one of four regimes: completely isolated; isolated but receiving a constant airflow from the mother colony; as previous but with the passage of nest volatiles towards the isolated ants blocked by adsorption on a SuperQ column; or reversed airflow direction-from the isolated ants to the nest interior. Ants that had been completely isolated for three weeks were subjected to aggressive behavior, but not those that had continued to receive airflow from the mother colony. Adsorbing the nest volatiles from the airflow by SuperQ abolished this difference, with these ants now also being subjected to aggression, indicating that nest volatiles can modulate nestmate recognition. Reverse airflow also reduced the level of aggression but to a lesser extent than airflow directed from the mother colony. In queenless colonies the overall aggression was reduced under all regimes, and there was no effect of flow, suggesting that the volatiles involved are queen-borne. The SuperQ adsorbed volatiles originated from Dufour's gland secretions of both workers and queen, implicating them in the process. Cuticular hydrocarbon profiles were not affected by exposure to nest volatiles, suggesting that the latter either constitute part of the recognition cues or affect worker behavior via a different, as yet elusive mechanism. © 2007 Elsevier Ltd. All rights reserved.Peer Reviewe