Here we report the genome sequence of the honeybee Apis mellifera, a key model for social behaviour and essential to global ecology through pollination. Compared with other sequenced insect genomes, the A. mellifera genome has high A+T and CpG contents, lacks major transposon families, evolves more slowly, and is more similar to vertebrates for circadian rhythm, RNA interference and DNA methylation genes, among others. Furthermore, A. mellifera has fewer genes for innate immunity, detoxification enzymes, cuticle-forming proteins and gustatory receptors, more genes for odorant receptors, and novel genes for nectar and pollen utilization, consistent with its ecology and social organization. Compared to Drosophila, genes in early developmental pathways differ in Apis, whereas similarities exist for functions that differ markedly, such as sex determination, brain function and behaviour. Population genetics suggests a novel African origin for the species A. mellifera and insights into whether Africanized bees spread throughout the New World via hybridization or displacement.

The evolution and maintenance of parthenogenetic species are a puzzling issue in evolutionary biology. Although the genetic mechanisms that act to restore diploidy are well studied, the underlying genes that cause the switch from sexual reproduction to parthenogenesis have not been analysed. There are several species that are polymorphic for sexual and parthenogenetic reproduction, which may have a genetic basis. We use the South African honeybee subspecies Apis mellifera capensis to analyse the genetic control of thelytoky (asexual production of female workers). Due to the caste system of honeybees, it is possible to establish classical backcrosses using sexually reproducing queens and drones of both arrhenotokous and thelytokous subspecies, and to score the frequency of parthenogenesis in the resulting workers. We found Mendelian segregation for thelytoky of egg-laying workers, which appears to be controlled by a single major gene (th). The segregation pattern indicates a recessive allele causing thelytoky. We found no evidence for maternal transmission of bacterial endosymbionts controlling parthenogenesis. Thelytokous parthenogenesis of honeybee workers appears to be a classical qualitative trait, because we did not observe mixed parthenogenesis (amphitoky), which might be expected in the case of multi-locus inheritance. Heredity (2005) 94, 533-537.

Differentiation into castes and reproductive division of labour are a characteristics of eusocial insects. Caste determination occurs at an early stage of larval development in social bees and is achieved via differential nutrition irrespective of the genotype. Workers are usually subordinate to the queen and altruistically refrain from reproduction. Workers of the Cape honeybee (Apis mellifera capensis) do not necessarily refrain from reproduction. They have the unique ability to produce female offspring parthenogenetically (thelytoky) and can develop into 'pseudoqueens'. Although these are morphologically workers, they develop a queen-like phenotype with respect to physiology and behaviour. Thelytoky is determined by a single gene (th) and we show that this gene also influences other traits related to the queen phenotype, including egg production and queen pheromone synthesis. Using 566 microsatellite markers, we mapped this gene to chromosome 13 and identified a candidate locus thelytoky, similar to grainy head (a transcription factor), which has been shown to be highly expressed in queens of eusocial insects. We therefore suggest that this gene is not only important for determining the pseudoqueen phenotype in A. m. capensis workers, but is also of general importance in regulating the gene cascades controlling reproduction and sterility in female social bees.

The innate immune system which helps individuals to combat pathogens comprises a set of genes representing four immune system pathways (Toll, Imd, JNK and JAK/STAT). There is a lack of immune genes in social insects (e.g. honeybees) when compared to Diptera. Potentially, this might be compensated by an advanced system of social immunity (synergistic action of several individuals). The bumble bee, Bombus terrestris, is a primitively eusocial species with an annual life cycle and colonies headed by a single queen. We used this key pollinator to study the temporal dynamics of immune system gene expression in response to wounding and bacterial challenge.Antimicrobial peptides (AMP) (abaecin, defensin 1, hymenoptaecin) were strongly up-regulated by wounding and bacterial challenge, the latter showing a higher impact on the gene expression level. Sterile wounding down-regulated TEP A, an effector gene of the JAK/STAT pathway, and bacterial infection influenced genes of the Imd (relish) and JNK pathway (basket). Relish was up-regulated within the first hour after bacterial challenge, but decreased strongly afterwards. AMP expression following wounding and bacterial challenge correlates with the expression pattern of relish whereas correlated expression with dorsal was absent. Although expression of AMPs was high, continuous bacterial growth was observed throughout the experiment.Here we demonstrate for the first time the temporal dynamics of immune system gene expression in a social insect. Wounding and bacterial challenge affected the innate immune system significantly. Induction of AMP expression due to wounding might comprise a pre-adaptation to accompanying bacterial infections. Compared with solitary species this social insect exhibits reduced immune system efficiency, as bacterial growth could not be inhibited. A negative feedback loop regulating the Imd-pathway is suggested. AMPs, the end product of the Imd-pathway, inhibited the up-regulation of the transcription factor relish, which is necessary for effector gene expression.

We present a set of 18 microsatellite DNA markers that can be run in two multiplex polymerase chain reactions as standard tool for assessing molecular ecological problems in honeybees (Apis mellifera). In addition to a set of six unlinked loci testing for classical population genetic parameters, we present three sets of four tightly linked loci, each located on three different chromosomes. These linked markers are useful for determining the number of colonies in a population as well as the parentage of drones and workers. Moreover, the tool kit can test for various modes of natural selection in honeybee populations.