The Evolution of Eusociality in Insects

Epigenetics in Social Insects: A New Direction for Understanding the Evolution of Castes

Originally written April, 2012 by Bryan White

Article 1 Source:

Epigenetics is a new field of biology that deals with an only recently discovered method of DNA inactivation called DNA methylation. DNA methylation is the process in which sections of DNA are methylated and primarily occurs on cytosines, although they could occur on any nucleotide. In this paper, the current state and understanding of DNA methylation and how it relates to the development and evolution of insect castes (particularly in the eusocial insect groups) is reviewed. Methylation is not the only possible epigenetic mechanism. DNA acetylation (the addition of acetyl molecules) is also possible, as well as ubiquitination (the addition of the ubiquitin protein).  However, DNA methylation is probably the most common. The end result of DNA methylation is the existence of a secondary language on top of the DNA language that can be modified by environmental factors, can be passed on to the next generation, and influence the development of offspring. DNA methylation can also have an evolutionary affect by increasing the rate of mutations in genes that are methylated for multiple generations, for genes that are inactivated can accumulate stop codons and other deleterious mutations. Based on this, the authors hypothesize that DNA methylation is potentially the primary method for caste selection in eusocial insects.

Epigenetics brings a whole new aspect to the table for understanding how castes evolved, and how castes are regulated (should a larva develop into a queen or worker?) in eusocial systems. In hymenopteran eusocial species, there is typically a vast amount of physical diversity amongst castes (workers, soldiers, queens and male drones), and workers have found it hard to explain this diversity using only genetic methods. This is largely due to the fact that it is well known that the development and selection of what a larva will develop into is environmentally based, but scientists do not have a clear idea of exactly how that developmental “decision” is enforced. Epigenetics stands as a good explanation for how environmental factors can influence larval development, and the authors suggest this probably carried out by the presence of DNA methylation genes such as DNMT3, coincidentally which Drosophila is lacking and so was thought unimportant. The direct connection between the expression of DNMT3 and the genes that are methylated is a new, expanding area of research.

Another one of the difficulties in understanding the evolution of eusociality has been trying to explain its evolution in terms of kin selection, specifically that haplodiploid species exhibit on average 75% more genetic relatedness of sisters than other species. The benefits of a haplodiploidy system as an example of kin selection theory were that it provided a strict means for both the regulation of sexual dimorphism (males are made up of only the queen’s genome) and suggested some involvement in the development of castes. However, epigenetics and DNA methylation offers a much better explanation for the existence of both large amounts of sexual dimorphism and phenotypic plasticity. DNA methylation has been found in many eusocial hymenopteran species, as well as primitively social hymenopterans, suggesting that DNA methylation is both a heavily conserved trait and is correlated to sociality, phenotypic plasticity and sexual dimorphism. Better understanding the phylogenetic location of insect groups that make use of DNA methylation can probably elucidate the question as to whether or not DNA methylation is the sole (or primary) source of caste determination.

The authors also attempt to lay out a conceptual framework for future studies, however I found their model unclear. What the authors seem to suggest is that eusociality is correlated with DNA methylation, but not a requirement. They do, however, do a good job outlining the specific areas of DNA methylation that need to be explored and understood to eliminate other possible explanations for the correlation between DNA methylation and eusociality, such as understanding the mechanistic effects that DNA methylation has on gene splicing and whether or not it is possible for eusocial insects to exhibit caste differentiation without DNA methylation genes.

Article 2 Source:

In this article the researchers hypothesize that up until this date, all progress on kin selection theory has largely been abstract in nature and not provided any concrete evidence for the theory. They argue that, in order for kin selection theory to be fulfilled in an empirical system, several stringent conditions must be met.

First, all interactions that are measured must be “additive and pairwise”, that is, they must only affect the pair of individuals involved in the interaction. This means that synergistic effects, such as the simultaneous cooperation of more than two individuals, are unable to be measured or incorporated into any mathematical model of kin selection.

Second, they argue that kin selection theory can only be applied to a very limited subset of population structures due to the requirement of global updating of interactions wherein global updating is the idea that any two individuals are competing uniformly for reproduction regardless of their geographic proximity to each other.

Third, they argue that if these two requirements are met, and they can only be met in some limited, artificial world, then when these requirements are met that the organismal interactions within that aforementioned world are also acting according to the conditions of natural selection theory, and that kin selection theory does not provide any additional biological information.

Finally, the authors also argue that the apparent simplicity of kin selection theory compared to that of natural selection theory is an illusion. Since the primary component of kin selection theory is the calculation of inclusive fitness, and the calculation of inclusive fitness requires the state of “all individuals whose fitness is affected by an action, not only those whose payoff is changed” to be known, then in effect kin selection theory is requiring the same information to be known as natural selection theory the state of all individuals affected rather than only those whose payoff (fitness) is increased.

In order to overcome the limitations imposed by kin selection theory, the authors propose a general, multi-level model of natural selection theory using only the general principals of population genetics. This model is used to explain how eusociality might evolve in five distinct evolutionary stages.

First, an organism must reach a state where there are clear groups within a population. Groups typically form around resources, nest sites, when parents and offspring stay together, or when flocks go to known breeding grounds.

Second, these groups begin to accumulate traits, otherwise known as pre-adaptations, that will increase the overall cohesion and cooperation of these groups. One such pre-adaptation is when a parent places large numbers of paralyzed prey around her eggs so that when the eggs hatch they will have a food source readily available, and then she moves on to create another nest. The next step towards eusociality would be for the parent to stay near the nest and guard the eggs until they are hatched. However, at this stage, the offspring will still leave the nest and so will the parent –  there is still dispersion.

Third is the evolution of clearly eusocial alleles, that is, traits that enforce the primary traits of eusociality. The key traits here are for individuals to stay in the nest instead of dispersing, and then other cooperative pre-adaptations can come into play.

Fourth is probably what can be called the optimization stage in which these eusocial alleles can be selected upon to reinforce the nest/colony structure.

Fifth is the final phase and selection now operates on the colonies instead of the individual organisms, and the evolution of more derived traits such as castes (workers/soldiers), fungal farming, aphid farming, and other highly cooperative activities. Here the authors have outlined the framework through which future studies can be conducted, most likely which will be a combination of behavioral ecology and phylogenetics. My criticisms of this paper can only be restricted to the authors’ use of the words “primitive” and “advanced”, which are common misnomers in evolutionary biology. A better term should be less derived or more derived, in reference to the ancestral state. For instance, the caste system of most ants is more derived compared to the loose grouping structure of some wasps.