The reactivity of neonatal rabbits to the mammary pheromone as a probe for viability

Newborn rabbits depend on a daily nursing interaction with the mother to gain milk and to survive. During this interaction, they localise and seize the nipples displaying a typical behaviour triggered by maternal odour cues. The mammary pheromone constitutes such a signal in domestic rabbits: it elicits sucking-related movements in more than 90% of the pups. However, some newborns remain unresponsive to the presentation of the pheromone, even pups apparently healthy and highly motivated to suck. The main goal of the present study was therefore to explore the link between the unresponsiveness of rabbit pups to the mammary pheromone and their growth and survival in breeding conditions. To that end, 293 newborns from 30 litters were tested for their head searching-oral grasping responses to the mammary pheromone on days 1 and 3, and their milk intake and mortality were followed up from days 1 to 21. It was hypothesised that unresponsive newborns would have subsequent difficulties in finding nipples, sucking and surviving. Early weight and success in milk intake were further considered as mediating factors in growth and viability. The results showed that pups that were unresponsive to the mammary pheromone on day 1 were less successful in gaining milk and had a higher rate of mortality than the responsive pups. However, this impact was modulated by the weight of pups: it appeared only in the lightest newborns. Moreover, this impact vanished on day 3. On the other hand, the pup weight and sucking success on days 1 to 3 strongly influenced viability and growth during the period extending from days 1 to 21. Taken together, the results show that the day-1 responsiveness of rabbit pups to the mammary pheromone can be considered as an indicator of individual viability in pups having a small weight (<48 g on day 1). The predictive validity of the pups’ pheromonal reactivity seems however time-limited as it works only during the first, but crucial, postnatal days.     

Electroantennogram responses of the Mediterranean fruit fly, Ceratitis capitata to identified volatile constituents from calling males

Fifty-six compounds from the odor of calling, sexually mature, laboratory reared males of the Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) were isolated by headspace trapping on Tenax columns and identified using GC/MS techniques (69 total compounds were detected). Electroantennogram responses (EAGs) to 54 of the 56 identified compounds as well as 5 analogs were tested on both sexes. Significant differences between the sexes in their responsiveness were found in 9 of the 54 identified compounds tested. There was no correlation between the amplitude of the EAG response and the relative abundance of compound identified from headspace analysis. Of the five major identified components, three elicited relatively small EAG responses, while two elicited large EAGs compared to the hexan-1-ol standard. The relative ranking of EAG responses were: methyl and ethyl hexenoates and hexanoates > C₄–C₆ esters and/or acetates > ethyl and methyl octenoates > monoterpenes > sesquiterpenes > C₂–C₅ acetates, alcohols and ketones. Behavioral bioassays on each of the five major identified components as well as a blend of six of the compounds showed some degree of attractancy to virgin females which in some cases approached the response to a pheromonal standard (male odors absorbed onto filter paper). These results are discussed in relationship to the insect's antennal sensitivity to putative pheromone components and/or allomonal components and to other reported C. capitata pheromone studies.

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Résumé Cinquante-six composés de l'odeur de mâles de C. capitata Weidemann, élevés en laboratoire, sexuellement mûrs et en appel, ont été isolés par piégeage sur colonnes tenax et identifiés par la technique GC/MS (69 composés avaient été détectés en tout). Les électroantennogrammes (EAGs) ont été examinés chez les deux sexes pour 54 des 56 composés identifiés et 5 de leurs analogues. Des différences significatives entre les sexes ont été observées pour 9 des 54 composés identifiés. Il n'y avait pas de corrélation entre l'ampleur de l'EAG et l'abondance relative du composé lors de son isolement. Pour les 5 principaux composés identifiés, 3 ont induit des EAGs relativement faibles, tandis que 2 étaient importants, par comparaison avec l'Hexane-1-ol utilisé comme témoin. Le classement relatif des EAG a été: hexénoates et hexanoates d'éthyl et de méthyl C₄–C₆ esters et/ou acétates octénoates d'éthyl ou de méthyl monoterpènes sesquiterpènes C₂–C₅ acétates, alcools et kétones. Les expériences de comportement avec chacun des 5 composés principaux identifiés, comme avec des mélanges de 6 composés ont mis en évidence une attraction des femelles vierges qui dans quelques cas avoisine la réponse à la phéromone témoin (odeur du mâle absorbée sur papier filtre). Ces résultats sont discutés en fonction de la sensibilité de l'antenne d'insexte aux composés supposés de la phéromone et aux composés allomonaux, et en fonction des autres études connues sur les phéromones de C. capitata.

     

Brood raiding and the population dynamics of founding and incipient colonies of the fire ant, Solenopsis invicta

Abstract. 1. One of the first activities of minim workers in incipient fire ant nests is mutual brood raiding, the amalgamation of nests through the reciprocal stealing of brood and defection of workers.
2. Discrete mating flights created cohorts of incipient colonies. About 25% of founding nests survived the claustral period of 16–48 days (depending on season). Early incipient colony mortality was 5–6% per day. Over 60% of this mortality was accounted for through brood raiding. Most colonies raided when each cohort first became active, and many raided more than once. Raid size (number of nests, duration, distance) tended to increase during the summer as colonies grew through raiding. After the raiding period, nest mortality rate dropped 3–10-fold. Only 1–3% of founding nests were still alive at this time.
3. Queens from losing or failed nests tended to abandon their nests and attempted to enter successful ones, often following raiding trails to do so. This emigration was at least as successful as non-emigration in ultimately achieving the status of reproductrix of a successful nest (about 4%).
4. Brood raiding is a dominant process in early population dynamics, probably accounting for most of the early nest mortality. Its effect is to change the venue and unit of competition from nest-against-nest to a shifting aggregation of queens, workers and brood involving entire local populations. Nest thinning is thus very rapid, and the boost to the size of winning nests very large, allowing raiding colonies to win the competition for territory, and to achieve the early colony maturity so important to this, and other, weedy species. The importance of winning brood raids may also have driven increased minim production through the evolution of pleometrosis.     

The behaviour of queen honeybees and their attendants

Abstract. The behaviour of queen and worker honeybees (Apis mellifera L.) was observed using small colonies in observation hives. Workers paid more attention to queens which had been mated for 2 months or more than to those which were newly mated; virgin queens received least attention. Queens received most attention when they were stationary and least when they were walking over the comb; virgin queens were most active. Queen cells had as many attendants as virgin queens and queen larvae were inspected almost continuously. The queen pheromone component 9–oxo-trans-2–decenoic acid stimulated 'court' behaviour when presented on small polyethylene blocks, but workers responded aggressively to complete extracts of queens' heads. Both the heads and abdomens of mated queens received much attention from court workers but the abdomens were palpated by more workers for longer and were licked much more. The queens' thoraces were least attended. Abdominal tergites posterior to tergite glands were licked for longer than those anterior to the glands. Only worker bees very near to the queen reacted to her and joined her 'court'.
No evidence was found of a diel periodicity in the behaviour of a queen or her 'court'. During the winter the queen's court was smaller than in summer and she walked less and laid fewer eggs. When colonies were fed with sucrose syrup in winter, their queens laid more eggs and workers reared more brood but there was no change in the attention received by the queens.
The implications of these findings for the secretion and distribution of queen pheromones are discussed.     

Queen succession in the primitively eusocial tropical waspRopalidia marginata (Lep.) (Hymenoptera: Vespidae)

Females of the primitively eusocial wasp Ropalidia marginata can be classified into three behavioral groups: Sitters, Fighters, and Foragers. It has been speculated that both Sitters and Fighters may be hopeful queens and that the Foragers may have little or no opportunities for direct reproduction. Here we show that in 9 of 12 queen-removal experiments where such a behavioral differentiation could be discerned, the individual that became a queen (the potential queen) was a Sitter in 6 cases, a Fighter in 2 cases, and a Forager in only 1 case. Although potential queens spent significantly more time absent from the nest and showed significantly higher rates of dominance behavior compared to the mean values for nonqueens in their colonies, they were intermediate with respect to all behaviors and age when compared to the range of values for nonqueens in their colonies. Potential queens were not necessarily the highest-ranking individuals among the nonqueens. The pattern of queen succession in this species appears to be quite different from the temperate pattern, where an old and active forager of high dominance rank is the potential queen. Although somewhat similar to the tropical pattern of a relatively younger female that has performed relatively little foraging being the potential queen, it is perhaps more accurate to describe the potential queens of R. marginata as unspecialized intermediates.     

Oviposition response of Simulium reptans (Diptera: Simuliidae) to the presence of conspecific eggs

Abstract.

• 1 The oviposition behaviour of Simulium reptans L. is described from two sites on the River Spey, Scotland. Female aggregations were observed immediately downstream of oviposition sites and were composed mainly of gravid flies (range 60–80%0) together with smaller numbers that were either freshly bloodfed (range 2–17%) or infected with mermithid nematodes (range 0–30%).
• 2 The time from landing on the oviposition sites to the onset of oviposition was recorded. The time in the presence of greater than 1-day-old eggs did not significantly differ from sites with no eggs present. However, the presence of freshly laid or I-day-old eggs significantly shortened the time to onset of oviposition. The cues that elicit oviposition are unknown but it is speculated that they may involve a pheromone.

     

Techniques for studying the execution of foreign queens by temporal subcastes in fire ants (Solenopsis invicta)

Using monogynous and polygynous colonies of the fire ant, Solenopsis invicta Buren, we developed methodology to determine which behavioural subcaste of workers was responsible for executing unfamiliar queens that were introduced to them. Separation of subcastes and introduction of queens to each of these separately was partly effective, but more accurate results were obtained by removing workers that were in the act of executing queens, marking them individually, and returning them to the colony. Their subsequent behaviour was then recorded to determine whether they behaved like nurses, reserves, or foragers. The results showed that foragers are dominant in execution behaviour and that all morphological subcastes (minors, medias, and majors) are involved. This methodology may be applied to other social insect species.

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Résumé A l'aide de colonies monogynes de S. invicta Buren, nous avons testé la sensibilité de deux méthodes: Premièrement, nous avons isolé les trois sous-castes temporelles des ouvrières (nurses, réserves et fourrageuses) auxquelles nous avons introduit des reines étrangères. Nous avons trouvé que les fourrageuses étaient le groupe le plus agressif, exécutant un pourcentage élevé des reines qui leur étaient présentées. Le degré de physogastrie de la reine étrangère, sa colonie d'origine et la présence de couvain n'avaient pas d'effet visible sur sa destinée. La sensibilité de cette méthode était limitée par la difficulté d'une séparation complète des souscastes temporelles.Deuxièmement, nous avons capturé des ouvrières exécutant activement des reines étrangères, après marquage nous les avons remises dans leur colonie pour observations ultérieures. Nous avons remarqué que leur localisation dans le nid et leur comportement correspondaient aux séquences trouvées pour les fourrageuses bien plus qu'à celles des ouvrières réserves ou nurses. Cette méthode est plus sensible. Des mesures de la largeur de la capsule céphalique de ces ouvrières ont montré qu'aucune des sous-caste morphologiques (mineurs, moyennes et majeurs) étaient dominantes dans le comportement d'exécution. Nos résultats indiquent que la sensibilité des ouvrières à certaines phéromones de la reine et odeur de la colonie augmente avec leur âge.

     

Social competition among gynes in halictine bees: The influence of bee size and pheromones on behavior

Solitary gynes of two species of social bees, Lasioglossum (Evylaeus) malachurumand L. (E.) pauxillum(Hymenoptera: Halictidae), were observed in the field and in the laboratory during the solitary (spring) phase of their life cycles. Fighting over nests among gynes of the former species is common when nests are being provisioned and can result in serious injury or even death to one or both interactants. The payoff is occasional acquisition of a nest. In contrast, fighting was never observed in the field among gynes of L. pauxillum.Several factors determine the outcome of such fights in L. malachuram,these include relative sizes of the two opponents as well as nest ownership. Dyadic interactions in the laboratory reveal that size influences the behavioral strategies of the gynes of both species, with the larger of two individuals in a dyadic interaction being on average more aggressive. Furthermore, in a second experimental series with L. malachurum,application to the smaller gyne of synthetically derived macrocyclic lactones found in the species' Dufour's gland pheromone mixture significantly decreased the aggressive tendencies of the larger such that its behavior was no longer significantly different from that for the smaller gyne. Therefore, among gynes, aggressive pheromonal signaling, coupled with other possible signal modalities, is probably an integral part in the communication system.     

Effects of social conditions on Juvenile Hormone mediated reproductive development in Bombus terrestris workers

Abstract. During the annual life cycle of the bumble bee Bombus terrestris (L.) colony, there is a stage characterized by worker reproduction in the presence of the queen. It has been proposed that this is a result of a decrease in queen inhibition. This hypothesis was examined by studying the effects of queens taken from colonies at different stages of development on several aspects of worker physiology and behaviour: rates of Juvenile Hormone (JH) release in vitro , ovary development, and behaviour associated with reproduction. After optimizing and validating the radiochemical assay for JH release for bumble bee workers, we found that queenless workers had significantly more developed ovaries and higher rates of release of JH than did queenright workers, confirming and extending previous findings that suggest that bumblebee ovarian development is under JH control. Mated queens, separated from their colony and brood, can have the same inhibitory effect on the reproductive development of callow workers. In contrast, workers confined with virgin queens or in queenless groups demonstrated a significantly higher rate of release of JH, overt aggression and threatening behaviours. However, there were no differences in rates of release of JH between workers confined in groups in the laboratory with queens taken from colonies either before or after the onset of worker reproduction. Furthermore, overt aggression and threatening behaviours were similar and low in both types of groups. These results gave no support to the hypothesis that a decrease in queen inhibition is associated with the onset of worker reproduction. We also show that young workers reared in colonies either before or after worker reproduction occurs, or in queenless colonies, all demonstrated similar, low rates of release of JH. These results suggest that older workers may inhibit the corpora allata of younger workers in queenless colonies.     

Ontogenetic patterns of volatiles identified in Dufour's gland extracts from queens and workers of the primitively eusocial halictine bee,Lasioglossum malachmum (Hymenoptera: Halictidae)

Summary Ontogenetic patterns of volatile compounds identified in Dufour's gland extracts from queens and workers of the primitively eusocial sweat beeLasioglossum malachurum (K.) were compared. Only young unmated queens showed high proportions of isopentenyl esters, while macrocyclic lactones were dominant in old breeding queens, spring queens, and workers. In young queens the relative and absolute amounts of volatiles changed one day after mating. A discriminant analysis revealed significant differences in odor patterns of unmated and mated young queens. The fat body was the largest in young females, while eggs could be recorded only in breeding queens. Possible functions of different odor components in the investigated female groups are discussed.