Ernesto Guzmán‐Novoa

One key advantage of eusociality is shared defense of the nest, brood, and stored food; nest defense plays an important role in the biology of eusocial bees. Recent studies on honey bees, Apis mellifera, have focused on the placement of defensive activity in the overall scheme of division of labor, showing that guard bees play a unique and important role in colony defense. Alarm pheromones function in integrating defensive responses; honey bee alarm pheromone is an excellent example of a multicomponent pheromonal blend. The genetic regulation of defensive behavior is now better understood from the mapping of quantitative trait loci (QTLs) associated with variation in defensiveness. Colony defense in other eusocial bees is less well understood, but enough information is available to provide interesting comparisons between A. mellifera and other species of Apis, as well as with allodapine, halictine, bombine, and meliponine bees. These comparative studies illustrate the wide variety of evolutionary solutions to problems in colony defense in the Apoidea.

A prominent theory states that animal phenotypes arise by evolutionary changes in gene regulation, but the extent to which this theory holds true for behavioral evolution is not known. Because "nature and nurture" are now understood to involve hereditary and environmental influences on gene expression, we studied whether environmental influences on a behavioral phenotype, i.e., aggression, could have evolved into inherited differences via changes in gene expression. Here, with microarray analysis of honey bees, we show that aggression-related genes with inherited patterns of brain expression are also environmentally regulated. There were expression differences in the brain for hundreds of genes between the highly aggressive Africanized honey bee compared with European honey bee (EHB) subspecies. Similar results were obtained for EHB in response to exposure to alarm pheromone (which provokes aggression) and when comparing old and young bees (aggressive tendencies increase with age). There was significant overlap of the gene lists generated from these three microarray experiments. Moreover, there was statistical enrichment of several of the same cis regulatory motifs in promoters of genes on all three gene lists. Aggression shows a remarkably robust brain molecular signature regardless of whether it occurs because of inherited, age-related, or environmental (social) factors. It appears that one element in the evolution of different degrees of aggressive behavior in honey bees involved changes in regulation of genes that mediate the response to alarm pheromone.

The first draft of the honey bee genome sequence and improved genetic maps are utilized to analyze a genome displaying 10 times higher levels of recombination (19 cM/Mb) than previously analyzed genomes of higher eukaryotes. The exceptionally high recombination rate is distributed genome-wide, but varies by two orders of magnitude. Analysis of chromosome, sequence, and gene parameters with respect to recombination showed that local recombination rate is associated with distance to the telomere, GC content, and the number of simple repeats as described for low-recombining genomes. Recombination rate does not decrease with chromosome size. On average 5.7 recombination events per chromosome pair per meiosis are found in the honey bee genome. This contrasts with a wide range of taxa that have a uniform recombination frequency of about 1.6 per chromosome pair. The excess of recombination activity does not support a mechanistic role of recombination in stabilizing pairs of homologous chromosome during chromosome pairing. Recombination rate is associated with gene size, suggesting that introns are larger in regions of low recombination and may improve the efficacy of selection in these regions. Very few transposons and no retrotransposons are present in the high-recombining genome. We propose evolutionary explanations for the exceptionally high genome-wide recombination rate.

This paper covers measures of field colony strength, by which we mean population measures of adult bees and brood. There are generally two contexts in which an investigator wishes to measure colony strength: 1. at the beginning of a study as part of manipulations to produce uniform colonies and reduce experimental error and; 2. as response variables during or at the end of an experiment. Moreover, there are two general modes for measuring colony strength: 1. an objective mode which uses empirical measures and; 2. a subjective mode that relies on visual estimates by one or more observers. There is a third emerging mode for measuring colony strength; 3. computer-assisted digital image analysis. A final section deals with parameters that do not directly measure colony strength yet give important indicators of colony state: flight activity at the entrance; comb construction; and two proxy measures of colony fitness: production of queen cells and drone brood. How to conduct different types of experiments are described, including infectious dose, dose effects, course of infection and longevity tests.