So far in this course we have dealt entirely with the evolution of characters that are controlled by simple Mendelian inheritance at a single locus. There are notes on the course website about gametic disequilibrium and how allele frequencies change at two loci simultaneously, but we didn’t discuss them. In every example we’ve considered we’ve imagined that we could understand something about evolution by examining the evolution of a single gene. That’s the domain of classical population genetics. For the next few weeks we’re going to be exploring a field that’s actually older than classical population genetics, although the approach we’ll be taking to it involves the use of population genetic machinery. If you know a little about the history of evolutionary biology, you may know that after the rediscovery of Mendel’s work in 1900 there was a heated debate between the “biometricians” (e.g., Galton and Pearson) and the “Mendelians” (e.g., de Vries, Correns, Bateson, and Morgan). Biometricians asserted that the really important variation in evolution didn’t follow Mendelian rules. Height, weight, skin color, and similar traits seemed to

Human biochemical genetics

I. the origin of species by means of natural selection

ION, GENERAL CONCEPTIONS, SELF-CONSCIOUSNESS, MENTAL INDIVIDUALITY. It would be very difficult for any one with even much more knowledge than I possess, to determine how far animals exhibit any traces of these high mental powers. This difficulty arises from the impossibility of judging what passes through the mind of an animal; and again, the fact that writers differ to a great extent in the meaning which they attribute to the above terms, causes a further difficulty. If one may judge from various articles which have been published lately, the greatest stress seems to be laid on the supposed entire absence in animals of the power of abstraction, or of forming general concepts. But when a dog sees another dog at a distance, it is often clear that he perceives that it is a dog in the abstract; for when he gets nearer his whole manner suddenly changes, if the other dog be a friend. A recent writer remarks, that in all such cases it is a pure assumption to assert that the mental act is not essentially of the same nature in the animal as in man. If either refers what he perceives with his senses to a mental concept, then so do both. (44. Mr. Hookham, in a letter to Prof. Max Muller, in the 'Birmingham News,' May, 1873.) When I say to my terrier, in an eager voice (and I have made the trial many times), "Hi, hi, where is it?" she at once takes it as a sign that something is to be hunted, and generally first looks quickly all around, and then rushes into the nearest thicket, to scent for any game, but finding nothing, she looks up into any neighbouring tree for a squirrel. Now do not these actions clearly shew that she had in her mind a general idea or concept that some animal is to be discovered and hunted? It may be freely admitted that no animal is self-conscious, if by this term it is implied, that he reflects on such points, as whence he comes or whither he will go, or what is life and death, and so forth. But how can we feel sure that an old dog with an excellent memory and some power of imagination, as shewn by his dreams, never reflects on his past pleasures or pains in the chase? And this would be a form of self-consciousness. On the other hand, as Buchner (45. 'Conferences sur la Theorie Darwinienne,' French translat. 1869, p. 132.) has remarked, how little can the hardworked wife of a degraded Australian savage, who uses very few abstract words, and cannot count above four, exert her self-consciousness, or reflect on the nature of her own existence. It is generally admitted, that the higher animals possess memory, attention, association, and even some imagination and reason. If these powers, which differ much in different animals, are capable of improvement, there seems no great improbability in more complex faculties, such as the higher forms of abstraction, and selfconsciousness, etc., having been evolved through the development and combination of the simpler ones. It has been urged against the views here maintained that it is impossible to say at what point in the ascending scale animals become capable of abstraction, etc.; but who can say at what age this occurs in our young children? We see at least that such powers

https://www.jstage.jst.go.jp/article/ase1887/22/257/22_257_458/_pdf

The descent of man and selection in relation to sex

In his introduction to this detailed survey of knowledge of insect societies, the author points out that research on insect sociology has proceeded in three phases, the natural history phase, the physiological phase and the population-biology phase. Advances in the first two phases have permitted embarkation in the third phase on a more rigorous theory of social evolution based on population genetics and writing this book, the author wished to relate the three phases of research on insects and to express insect sociology as population biology. A glossary of terms, a considerable bibliography and a general index are included. Other CABI sites 

The Insect Societies

Meta-analytic procedures for social research

The evolution of group living requires that individuals limit the inherent risks of parasite infection. To this end, group living insects have developed a unique capability of mounting collective anti-parasite defences, such as allogrooming and corpse removal from the nest. Over the last 20 years, this phenomenon (called social immunity) was mostly studied in eusocial insects, with results emphasizing its importance in derived social systems. However, the role of social immunity in the early evolution of group living remains unclear. Here, I investigate this topic by first presenting the definitions of social immunity and discussing their applications across social systems. I then provide an up-to-date appraisal of the collective and individual mechanisms of social immunity described in eusocial insects and show that they have counterparts in non-eusocial species and even solitary species. Finally, I review evidence demonstrating that the increased risks of parasite infection in group living species may both decrease and increase the level of personal immunity, and discuss how the expression of social immunity could drive these opposite effects. By highlighting similarities and differences of social immunity across social systems, this review emphasizes the potential importance of this phenomenon in the early evolution of the multiple forms of group living in insects.

https://royalsocietypublishing.org/doi/pdf/10.1098/rstb.2014.0102

Social immunity and the evolution of group living in insects

Parental care increases the fitness of offspring at a cost to the parents in terms of residual reproductive success. This trade-off may be affected by ecology, life history and the social environment, which raises the question as to how these factors contribute to the evolution of parental care. Here, previous hypotheses concerning the evolution of parental care in insects are summarized and discussed and the underlying empirical evidence is reviewed.
Ecological factors such as harsh environments, ephemeral food sources or predation pressure are broadly accepted as evolutionary drivers of parental care. The most consistent evidence supports a role for natural enemies such as predators, microbes and cannibalistic conspecifics. Also, the importance of ecological factors may interact with the life history (parity) of a species, either as a pre-adaptation facilitating the evolution of parental care or as a consequence of enhanced parental investment under parental care. Yet, only limited experimental research has been carried out to test the combined influence of ecology and life history in the evolution of parental care.
Several forms of care can mediate the transition from solitary to family living, which entails the emergence of a novel – social – environment that generates new selection pressures from interactions within and between families. In this context, we review examples of studies on communal breeding, brood parasitism, parent–offspring conflict and co-adaptation, and discuss how these social interactions may in turn be influenced by ecological factors such as food availability or population density.
Insects are uniquely suitable for experimental and comparative research on the complex interplay between ecology, life history, and the social environment.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/een.12000

The evolution of parental care in insects: the roles of ecology, life history and the social environment

Many insects raise their offspring on organic substrates or in the soil where microorganisms are abundant. Microbes may pose a serious threat to offspring development and survival by either decomposing food resources or directly infecting the offspring. Selection to cope with these effects may favor social defenses, for example, through forms of parental care that can limit or eliminate these threats to offspring fitness. In this study, we experimentally tested if maternal egg attendance in the European earwig Forficula auricularia has a function as a social defense against mold infection of eggs by manipulating exposure of eggs to mold spores and the presence of the mother in a fully factorial design. Furthermore, we investigated the potential roles of egg grooming behavior and maternal transfer of chemicals as underlying mechanisms. As predicted, the beneficial effect of egg attendance on hatching success was significantly enhanced when eggs were exposed to the mold. Females significantly increased their egg grooming duration in response to mold exposure of her eggs, and the quantity of chemicals (identified as hydrocarbons) was maintained among attended eggs but decreased substantially among unattended eggs. Maternal transfer of chemicals was confirmed in extractions of glass beads that were mingled into attended or unattended clutches. This study shows that maternal egg attendance in the European earwig has a social defense function protecting offspring against mold infection. The maternal egg grooming behavior seems to be key for this effect, probably through both the mechanical removal of spores and the continued application of chemical substances on the egg surface.

Maternal care provides antifungal protection to eggs in the European earwig

The study of sex allocation in social Hymenoptera (ants, bees, and wasps) provides an excellent opportunity for testing kin-selection theory and studying conflict resolution. A queen-worker conflict over sex allocation is expected because workers are more related to sisters than to brothers, whereas queens are equally related to daughters and sons. If workers fully control sex allocation, split sex ratio theory predicts that colonies with relatively high or low relatedness asymmetry (the relatedness of workers to females divided by the relatedness of workers to males) should specialize in females or males, respectively. We performed a meta-analysis to assess the magnitude of adaptive sex allocation biasing by workers and degree of support for split sex ratio theory in the social Hymenoptera. Overall, variation in relatedness asymmetry (due to mate number or queen replacement) and variation in queen number (which also affects relatedness asymmetry in some conditions) explained 20.9% and 5% of the variance in sex allocation among colonies, respectively. These results show that workers often bias colony sex allocation in their favor as predicted by split sex ratio theory, even if their control is incomplete and a large part of the variation among colonies has other causes. The explanatory power of split sex ratio theory was close to that of local mate competition and local resource competition in the few species of social Hymenoptera where these factors apply. Hence, three of the most successful theories explaining quantitative variation in sex allocation are based on kin selection.

/pdf/split-sex-ratios-in-the-social-hymenoptera-a-meta-analysis-44oltqk5d4.pdf

Split sex ratios in the social Hymenoptera: a meta-analysis

Whether to reproduce once or multiple times (semelparity vs. iteroparity) is a major life-history decision that organisms have to take. Mode of parity is usually considered a species characteristic. However, recent models suggested that population properties or condition-dependent fitness payoffs could help to maintain both life-history tactics within populations. In arthropods, semelparity was also hypothesised to be a critical pre-adaptation for the evolution of maternal care, semelparous females being predicted to provide more care due to the absence of costs on future reproduction. The aim of this study was to characterize potential fitness payoffs and levels of maternal care in semel- and itero-parous females of the European earwig Forficula auricularia. Based on 15 traits measured in 494 females and their nymphs, our results revealed that iteroparous females laid their first clutch earlier, had more eggs in their first clutch, gained more weight during the 2 weeks following hatching of the first clutch, but produced eggs that developed more slowly than semelparous females. Among iteroparous females, the sizes of first and second clutches were significantly and positively correlated, indicating no investment trade-off between reproductive events. Iteroparous females also provided more food than semelparous ones, a result contrasting with predictions that iteroparity is incompatible with the evolution of maternal care. Finally, a controlled breeding experiment reported full mating compatibility among offspring from females of the two modes of parity, confirming that both types of females belong to one single species. Overall, these results indicate that alternative modes of parity represent coexisting life-history tactics that are likely to be condition-dependent and associated with offspring development and specific levels of maternal care in earwigs.

/pdf/one-clutch-or-two-clutches-fitness-correlates-of-coexisting-27megp39ji.pdf

Joël Meunier

Papers

Social immunity and the evolution of group living in insects

The evolution of parental care in insects: the roles of ecology, life history and the social environment

Maternal care provides antifungal protection to eggs in the European earwig

Split sex ratios in the social Hymenoptera: a meta-analysis

One clutch or two clutches? : Fitness correlates of coexisting alternative female life-histories in the European earwig