On the relationship between aggression and reproduction in pairs of orphaned worker bumblebees (Bombus impatiens)

This study characterized aggression and reproduction within pairs of orphaned bumblebee sisters (Bombus impatiens (Cresson, 1863)). Twenty-one pairs were filmed in the laboratory over 5–10 days. Frequencies of aggression and egg-laying were obtained for each bee, and the presence or absence of brood was manipulated. Aggression and egg-laying were more likely to co-occur in pairs placed without brood compared to pairs placed with brood. A significant positive correlation was found between members of a pair in the rates of aggression. In addition, a strong positive correlation was found in their rates of egg-laying: bees that had more sons also tended to have more nephews. The results show that under conditions of unrestricted food availability, behavioural interactions are compatible with continued reproduction by both orphaned workers. Though aggression may limit reproduction, it seems either to be an ineffective means of obtaining a reproductive monopoly in some situations and/or to be a set of behaviours invested with other possible functions.     

A conceptual model for the origin of worker behaviour and adaptation of eusociality

In a model based on the wasp family Vespidae, the origin of worker behaviour, which constitutes the eusociality threshold, is not based on relatedness, therefore the origin of eusociality does not depend on inclusive fitness, and workers at the eusociality threshold are not altruistic. Instead, incipient workers and queens behave selfishly and are subject to direct natural selection. Beyond the eusociality threshold, relatedness enables 'soft inheritance' as the framework for initial adaptations of eusociality. At the threshold of irreversibility, queen and worker castes become fixed in advanced eusociality. Transitions from solitary to facultative, facultative to primitive, and primitive to advanced eusociality occur via exaptation, phenotypic accommodation and genetic assimilation. Multilevel selection characterizes the solitary to highly eusocial transition, but components of multilevel selection vary across levels of eusociality. Roles of behavioural flexibility and developmental plasticity in the evolutionary process equal or exceed those of genotype.     

Parasites and sexual reproduction in psychid moths

Persistence of sexual reproduction among coexisting asexual competitors has been a major paradox in evolutionary biology. The number of empirical studies is still very limited, as few systems with coexisting sexual and strictly asexual lineages have been found. We studied the ecological mechanisms behind the simultaneous coexistence of a sexually and an asexually reproducing closely related species of psychid moth in Central Finland between 1999 and 2001. The two species compete for the same resources and are often infected by the same hymenopteran parasitoids. They are extremely morphologically and behaviorally similar and can be separated only by their reproductive strategy (sexual vs. asexual) or by genetic markers. We compared the life-history traits of these species in two locations where they coexist to test predictions of the cost-of-sex hypothesis. We did not find any difference in female size, number of larvae, or offspring survival between the sexuals and asexuals, indicating that sexuals are subject to cost of sex. We also used genetic markers to check and exclude the possibility of Wolbachia bacteria infection inducing parthenogenesis. None of the samples was infected by Wolbachia and, thus, it is unlikely that these bacteria could affect our results. We sampled 38 locations to study the prevalence of parasitoids and the moths' reproductive strategy. We found a strong positive correlation between prevalence of sexual reproduction and prevalence of parasitoids. In locations where parasitoids are rare asexuals exist in high densities, whereas in locations with a high parasitoid load the sexual species was dominant. Spatial distribution alone does not explain the results. We suggest that the parasite hypothesis for sex may offer an explanation for the persistence of sexual moths in this system.     

Evidence for competition between honeybees and bumblebees; effects on bumblebee worker size

Numerous studies suggest that honeybees may compete with native pollinators where introduced as non-native insects. Here we examine evidence for competition between honeybees and four bumblebee species in Scotland, a region that may be within the natural range of honeybees, but where domestication greatly increases the honeybee population. We examined mean thorax widths (a reliable measure of body size) of workers of Bombus pascuorum, B. lucorum, B. lapidarius and B. terrestris at sites with and without honeybees. Workers of all four species were significantly smaller in areas with honeybees. We suggest that reduced worker size is likely to have implications for bumblebee colony success. These results imply that, for conservation purposes, some restrictions should be considered with regard to placing honeybee hives in or near areas where populations of rare bumblebee species persist.     

Queen influence on worker reproduction in bumblebees (<Emphasis Type="Italic">Bombus terrestris</Emphasis>) colonies

Summary The bumblebee Bombus terrestris is a good model in which to study the regulation of worker reproduction, because the onset of queen-worker conflicts regarding male production is constrained by the annual life cycle of the colony. Worker reproduction in this species is inhibited until late in colony development. The underlying proximate mechanism suggested for this delayed worker reproduction is queen control using combined pheromonal and behavioural determinants. The volatile queen pheromone hypothesis was tested by monitoring the length of time necessary for worker reproduction to occur when workers were separated from the queen by a double screen. These workers always reproduced before the onset of the competition phase in the queenright compartment (QRC), after the time lag required for egg maturation. Since the double-mesh-separated compartment was genuinely a queenless compartment, the volatile queen pheromone hypothesis was refuted.The possible involvement of a non-volatile pheromone and/or of physical intimidation by the queen was then tested using a queen excluder, which allowed the workers, but not the queen, to travel freely between two nest compartments, creating a refuge from the queen. Although worker reproduction in the queen-excluded compartment (QEC) was delayed compared to the queenless situation, it still occurred before the onset of the competition phase in the QRC and after the onset of queen production.These results indicate that workers autoregulate their reproduction rather than it being controlled by the queen (pheromonal or behavioural). The possible factors affecting this autoregulation may be 1) the workers perception of the queen and, possibly, her status through pheromonal emission and 2) the workers perception that the larvae are committed to become queens.Received 18 November 2003; revised 24 February 2004; accepted 1 March 2004.     

The genetic architecture of immune defense and reproduction in male Bombus terrestris bumblebees

Understanding the architecture of genetic variation, that is the number, effect, location, and interaction, of genes responsible for phenotypic variability in nature is important for the understanding of microevolutionary processes. In this study, we have used a quantitative trait loci (QTL) approach to uncover the genetic architecture of fitness-relevant traits associated with reproduction and immune defense in male Bombus terrestris bumblebees. Three male reproductive investment traits, the number and length of the produced sperm and the size of the accessory glands, were studied. Two branches of the insect innate immune system, the activation of the Phenoloxidase-cascade and the hemolymph's antibacterial activity, were investigated. We found that variation in most of the studied traits is based on a network of minor QTLs and epistatic interactions. Unexpectedly, there was no evidence for phenotypic or genetic trade-offs between the presumably costly investment in immune defense and reproductive effort in this population for the measured traits. In fact, we found a positive correlation, both, in phenotype and genetic architecture for the number of produced sperm and antibacterial activity against an insect pathogen. A major finding for all traits analyzed was that the epistatic interactions accounted for a major proportion of the explained phenotypic variance. Especially for traits involved in immune defense, this pattern highlights the possible role of parasites in the evolution and maintenance of recombination and sexual reproduction.     

Effects of selection for honey bee worker reproduction on foraging traits

The “reproductive ground plan” hypothesis (RGPH) proposes that reproductive division of labour in social insects had its antecedents in the ancient gene regulatory networks that evolved to regulate the foraging and reproductive phases of their solitary ancestors. Thus, queens express traits that are characteristic of the reproductive phase of solitary insects, whereas workers express traits characteristic of the foraging phase. The RGPH has also been extended to help understand the regulation of age polyethism within the worker caste and more recently to explain differences in the foraging specialisations of individual honey bee workers. Foragers that specialise in collecting proteinaceous pollen are hypothesised to have higher reproductive potential than individuals that preferentially forage for nectar because genes that were ancestrally associated with the reproductive phase are active. We investigated the links between honey bee worker foraging behaviour and reproductive traits by comparing the foraging preferences of a line of workers that has been selected for high rates of worker reproduction with the preferences of wild-type bees. We show that while selection for reproductive behaviour in workers has not altered foraging preferences, the age at onset of foraging of our selected line has been increased. Our findings therefore support the hypothesis that age polyethism is related to the reproductive ground plan, but they cast doubt on recent suggestions that foraging preferences and reproductive traits are pleiotropically linked.     

The consequences of earlier reproduction in declining populations

Conflicting claims have been published concerning the consequences of earlier reproduction. In particular, it has been claimed that earlier reproduction will increase the rate of decline in a declining population, have no effect in a stable population, and enhance the rate of growth in an increasing population. However, if fecundity and mortality are conceptually separated so that earlier reproduction is defined as moving the fecundity column of the life table forward in time but not changing the mortality column, then earlier reproduction will nearly always cause the rate of change of a population to become more positive. This definition is believed to be more consistent with what biologists think of as earlier reproduction. Also, this concept requires that the cost of reproducing earlier be explicitly addressed rather than hiding it in assumptions implicit in a definition of earlier reproduction. This paper reconciles published mathematical demography models with published biological claims concerning earlier reproduction.     

Colony fusion and worker reproduction after queen loss in army ants

Theory predicts that altruism is only evolutionarily stable if it is preferentially directed towards relatives, so that any such behaviour towards seemingly unrelated individuals requires scrutiny. Queenless army ant colonies, which have anecdotally been reported to fuse with queenright foreign colonies, are such an enigmatic case. Here we combine experimental queen removal with population genetics and cuticular chemistry analyses to show that colonies of the African army ant Dorylus molestus frequently merge with neighbouring colonies after queen loss. Merging colonies often have no direct co-ancestry, but are on average probably distantly related because of overall population viscosity. The alternative of male production by orphaned workers appears to be so inefficient that residual inclusive fitness of orphaned workers might be maximized by indiscriminately merging with neighbouring colonies to increase their reproductive success. We show that worker chemical recognition profiles remain similar after queen loss, but rapidly change into a mixed colony Gestalt odour after fusion, consistent with indiscriminate acceptance of alien workers that are no longer aggressive. We hypothesize that colony fusion after queen loss might be more widespread, especially in spatially structured populations of social insects where worker reproduction is not profitable.     

The consequences for human reproduction of a robust inflammatory response

Innate and adaptive immune responsiveness is variable within the population. Since robust immune reactions are critical to the survival of humans, the existence of immune variability in the population suggests the existence of competing, alternative phenotypes. Although women with powerful immune responsiveness may be more likely to survive to reproduce, their reproductive experiences may be less successful than women who are not as responsive. Normal pregnancy elicits a maternal inflammatory reaction. This can be understood on the basis of maternal-fetal conflict theory: inflammation is a component of the maternal attempt to limit excessive fetal demands. However, an overly aggressive inflammatory reaction has been shown to relate to a variety of adverse reproductive outcomes. Reviewed here are several examples, including the fallopian tube damage that results from pelvic inflammatory disease, the upregulated inflammatory response among women who develop preeclampsia, an association between immune hyperresponsiveness and premature delivery, and the relationship between autoimmune diseases and multiple adverse pregnancy outcomes. The hypothesis that immune hyperresponsiveness limits reproductive capacity suggests many avenues for research.     

Chemical correlates of reproduction and worker policing in a myrmicine ant

In a number of wasps, bees, and in particular ponerine ants, quantitative and qualitative variation in the profile of cuticular hydrocarbons is associated with variation in fecundity and is likely to serve for communicating the reproductive status of an individual. Here, we demonstrate that the chemical profile on the cuticle of fertile workers and queens of the myrmicine ant Temnothorax unifasciatus is different from that of non-reproductive workers. Fertility and apparently also cuticular signatures are reversible under the influence of policing by worker aggression. Though no policing by egg-eating occurs in this species, queen and worker laid eggs also differed in their chemical profile.     

Aspects of worker reproduction in four social wasp species (Insecta: Hymenoptera: Vespidae)

The question of male production by workers in social wasp colonies (Vespinae) has long concerned researchers. Its occurrence would suggest that queen control of colony reproduction is not absolute and that the evolution and maintenance of eusociality in vespines may have been promoted through direct reproductive ability of the worker caste. Dissections of over 3,300 workers from 34 Paravespula and Vespula colonies failed to reveal a significant contribution to male production by workers in most colonies. In those colonies which did contain ovary-developed workers, the wasps tended to be larger than their nestmates in two of the species. On the other hand, ovary-developed workers could not be characterized as specializing in intra- or extranidal tasks in any species. The onset of worker ovarian development was associated with increased size of nests and occurred predominantly in late-season colonies. The near-complete sterility of workers through most of the season suggests that models invoking mutualism and characterizing workers as 'hopeful reproductives' cannot explain the maintenance of eusociality in vespine wasps.     

Towards greater realism in inclusive fitness models: the case of worker reproduction in insect societies

The conflicts over sex allocation and male production in insect societies have long served as an important test bed for Hamilton''s theory of inclusive fitness, but have for the most part been considered separately. Here, we develop new coevolutionary models to examine the interaction between these two conflicts and demonstrate that sex ratio and colony productivity costs of worker reproduction can lead to vastly different outcomes even in species that show no variation in their relatedness structure. Empirical data on worker-produced males in eight species of Melipona bees support the predictions from a model that takes into account the demographic details of colony growth and reproduction. Overall, these models contribute significantly to explaining behavioural variation that previous theories could not account for.     

Degrees of disruption: projected temperature increase has catastrophic consequences for reproduction in a tropical ectotherm

Although climate change models predict relatively modest increases in temperature in the tropics by the end of the century, recent analyses identify tropical ectotherms as the organisms most at risk from climate warming. Because metabolic rate in ectotherms increases exponentially with temperature, even a small rise in temperature poses a physiological threat to tropical ectotherms inhabiting an already hot environment. If correct, the metabolic theory of climate warming has profound implications for global biodiversity, since tropical insects and arachnids constitute the vast majority of animal species. Predicting how climate change will translate into fitness consequences for tropical arthropods requires an understanding of the effects of temperature increase on the entire life history of the species. Here, in a comprehensive case study of the fitness consequences of the projected temperature increase for the tropics, we conducted a split‐brood experiment on the neotropical pseudoscorpion, Cordylochernes scorpioides, in which 792 offspring from 33 females were randomly assigned at birth to control‐ and high‐temperature treatments for rearing through the adult stage. The diurnally varying, control treatment temperature was determined from long‐term, average daily temperature minima and maxima in the pseudoscorpion's native habitat. In the high temperature treatment, increasing temperature by the 3.5 °C predicted for the tropics significantly reduced survival and accelerated development at the cost of reduced adult size and a dramatic decrease in level of sexual dimorphism. The most striking effects, however, involved reproductive traits. Reared at high temperature, males produced 45% as many sperm as control males, and females failed to reproduce. Sequencing of the mitochondrial ND2 gene revealed two highly divergent haplogroups that differed substantially in developmental rate and survivorship but not in reproductive response to high temperature. Our findings suggest that reproduction may be the Achilles’ heel of tropical ectotherms, as climate warming subjects them to an increasingly adverse thermal environment.