Much of the scientific study of human social behaviour — human behavioural ecology, sociobiology, and its various associates — owes a great deal to the study of the insect world. There is of course much to readily criticise such for; humans and insects are very diverse forms of life, much of their physiology and life history quite alien to the other. Indeed, because of a matter of scale — their size — it is not unreasonable to assert insects occupy a world apart from ours. More properly: it is a world within our own, a world that exists within the indeterminable nooks and crannies of our own place of inhabitance, teeming within the countless cracks and crevices of all the places on earth except for its deepest, darkest oceanic trenches, the bittermost cold of the polar reaches, and the intolerable, molten heats of the earth’s depths. Even so, it is a world that is far more expansive than what we know. We live on a thin surface, the edges of a heavily-folded, ecological sheet, our size, tolerances, and senses forbidding us from personally squeezing our way down in to its prohibitive folds, where only little, extraordinary things, of diverse shapes and incredible abilities, may slip into. Like the figurative sheet, once it is stretched out — dissecting and spreading open the countless secret passages through soil, wood, and skies — its true surface is revealed to be an area of unchartered vastness, an infinitude of microcosms among which the occupancy of man winds about as single thread.
One should always be cautious about applying what is or what works for one kind of these denizens of bizarre habits and habitats to what is — the biophysical rules — that dictate the limitations of another kind. This is, generously, something of the argument grasped and thrust forth by those keen to refute any and all of the behavioural and biological maxims set forth by a variety of sciences, including the aforementioned sociobiology and HBE. Especially when the subject of scientific scrutiny is humanity — we often like to assert the cherished modern notion of every individual being irreducible to a set of seemingly-cold, mechanical principles (or, more likely the ticket, the odiousness lies in humans being set besides, comparable to other species, the abhorred “beasts” classically villified by human exceptionalism). The irony is not lost in that most such “champions of individuality,” the irreducible nature of the human character, will happily be reduced to a set of ordinal categories for demographic statistics and the homogenising, reductionist, official policies of the State formed out of such — suppositions often far cruder, less-critical, and with more punitive, “dehumanising” fruits than assertions set forth in the sociobiological literature. A chorus of such criticisms from these part-time champions inaugurated the publishing of Sociobiology: the New Synthesis (1975), written by the acclaimed E.O. Wilson — a myrmecologist, “ant-ecologist,” a student of the insect world. Then as now, the contesting clamour was largely aimed at his single chapter on “Man,” which briefly held the same sociobiological lens used to examine other species over mankind. For its brevity in handling this one specimen, one might not have anticipated the large and sustained backlash, to it and the namesake field it founded, from a wide range of seemingly-disparate parties. Religious groups, “conservative” representatives, and liberals alike have all found reason to be leery of the work when exposed to its implications (which is not to say there aren’t religious, conservative, or liberal individuals who have praised it or regarded it neutrally) — what unites them is, again, a strong attachment to the common, sentimental denominator of modern civilisation, that of human exceptionalism and our supposed, crowning achievement — modern civilisation and “progress” — and the abhorrence of what is “natural,” “primitive,” “beastly,” “backwards,” from which the “civil” man must distinguish himself.
That was to give a vignette of much of the popular opposition to the endeavours of sociobiology and its derivative of HBE. The less-popular opposition, largely reserved to scientific discourse, will often come around to the precept set forth already — the problems of comparing one form of life to another. But for all of the virtues of this sceptical glance, one must acknowledge that in all of its rich diversity, life is yet life. It emerged from a common source eons ago, elaborating upon itself, branching out and bursting forth into flowers of various shape and perfumes, embellishing itself by slight degrees with every passing generation, if the environment that nurtures it so demands or allows. From these common ancestors we received and hence share the most basic, the most foundational of the building blocks of life: self-replicating molecules — RNA, DNA — that, through the magic of a few acids, sequesters and recruits a variety of other substances and electric energy itself to recreate itself, to reinvent itself by navigating, feeling, responding to its environment. And the environment, physical reality itself — shaped and formed on our world by life to an extraordinarily complex degree — is a commonality we share as well, each of us subject to many of the same physical laws, to which our physiological or behavioural responses may diverge or converge in accordance to our inherited physical tolerances. In short, all life is related, by virtue of evolving in a shared reality, by which the many worlds — our own, or the ones belonging to various insect life, for example — are but the interlocked weave of the tapestry of reality. We would as such be able to rightfully identify limitations and abilities of various similitudes between even the furthest flung members of life, such as ourselves and the dwellers of the deeper places. And so the value of studying this world within and surrounding our own, the world of the insects — for “as above, so below,” that when we look upon the small, we will always look on and learn something of ourselves, however slight, for we look upon a relative who trod a different path from ourselves long ago, a creature made of similar stuffs, however bizarre the shape, passing through the same air and over the same earth, even if the parts they make use of is narrower or wider than our own niche. The differences of course cannot be dismissed; but neither can the things we share in common, and the generalisations and relations we might discern from it.
With that in mind, I would like to illustrate one possible relation, one lesson to be learned by gazing into the Below of the Insecta. Just to clarify, these are my own thoughts on the matter, a philosophical extrapolation from a scientific study whose authors may never have thought of it in any such light or who may disagree with what I have to say. Like much of the lessons I take away from interactions with or contemplation of others, be they human or some other lifeform, I have tried to relate it to a larger picture of the world, how it functions — and, again, what is reflected of oneself and one’s state in these deep and strange pools of life. The study in question was detailed in an article by Kapheim et al. (2015), entitled “Kinship, Parental Manipulation and Evolutionary Origin of Eusociality,” and, as the name indicates, it examined how the phenomena of eusociality might have evolved throughout the natural world (using its classic insect models, the Hymenopterans, i.e. “wasps,” “bees,” and “ants”). Just to clarify in case that word/concept of “eusociality” is unfamiliar to you: it is a form of sociality amongst animals in which conspecifics engage in cooperative breeding by having a division of labour and reproductive castes. Ants are often the usual example demonstrating the phenomena: most members of a society/colony cannot reproduce, being sterile females who instead will work on either guarding the nest, taking care of the brood, foraging, etc., while there is a much smaller number of reproductives who will create more workers and the handful of fertile individuals (the “queen” and any males involved) to continue the colony or go off to establish their own.
Figure 1: Diagram of a generalised life cycle and castes of termites, another eusocial insect. They are not related to ants/wasps, being instead in the same taxonomic order as cockroaches. A nymph (immature insect) can either develop into a worker, soldier, or a reproductive individual. In some termite species, there is no discernible “worker caste,” the workers instead remaining nymphs/immatures. Some reproductives, known as “alates,” have wings so that they can leave their colony, disperse, and potentially be mated to establish their own colony.
The usual assumption of how such a thing would evolve is that it is a kind of extreme inclusive fitness via kin selection. With ants/wasps/bees in particular, the way their genetic material is distributed amongst offspring results in female siblings who are supposed to be 75% related to each other. This system of genetic inheritance, known as “haplodiploidy,” is quite foreign to what you will find amongst most mammals, humanity included, in which siblings are typically only 50% related to one another. By cooperating with their sisters to protect and help their mother, female Hymenopterans could then pass on 75% of their genetic material by their mother’s continued successful reproduction of more sisters for them, some of whom will be reproductive, without having to reproduce themselves. Since having your own offspring/brood is no easy feat due to the time and energy investment demanded, it is then supposed that natural selection through kin selection could act in such a way that eusocial organisations can eventually emerge by selecting for individuals who are increasingly cooperative with their sisters and their mother, as this cooperation allows for greater fitness benefits to them than the individual would get from going out and directly reproducing themselves, founding their own nest/laying their own eggs and incurring all the costs and risks associated with that. This then assumes that the evolution of eusociality is driven by worker “altruism,” and these proposed benefits of helping one’s “kin” to pass on some of your genes embodied in them is “inclusive fitness” by kin selection.
However, there are a number of problems with such a supposition, one of them being that not all the workers are actually that closely related (75%). If the queen mates more than once, with a different male, then that brood will not be as closely related to the existing members. And yet they still they cooperate, even though there is clearly not as great a fitness benefit there as supposed for evolution to act upon. With that being the case, i.e. most of the sisters you are helping, and the few reproductives who will carry on into the future, are only as closely related to you as you would find among non-social/non-eusoscial mammalian siblings, it becomes very questionable how much individuals are really benefitting in terms of fitness from such a system, or how this alone could drive a species to becomes eusocial. To which, there is an alternative explanation for the evolution of eusociality — that of “parental parasitism” or “maternal manipulation.” As the name implies, this theory supposes that there really isn’t much of a fitness benefit for the workers/sterile individuals to forgo their own reproduction and help the reproductive elites, at least not compared to reproducing oneself. Instead, the parents or the mother/queen merely manipulate the offspring into helping them, drastically improving their own fitness, irrespective of the fitness cost or benefit to the workers.
This manipulation can be done genetically, epigenetically, or by social/chemical cues. In honey bees, for example, workers are created by feeding the larvae an inferior food, “bee bread,” while reproductive queens are created by feeding the larvae a richer food known as “royal jelly.” Queens of certain Hymenopteran species can also exude pheromones that delay or prevent the offspring from maturing and reproducing, such that they will continue helping her instead of attempting to create their own brood. One might entertain the idea that “primitively-social” bumblebees represent an earlier, “incomplete” offshoot to a eusocial track. The queen will form a colony, but she does not have perfect control over her daughter’s reproductive abilities, such that daughters may begin to try to lay their own eggs at some point. The queen, however, will go about destroying them, until, eventually, late in the season, the colony undergoes a collapse, in which the daughters/workers eventually turn on the queen and kill her, though at that point, it is too late for the daughters to establish any viable colony of their own and the reproductive offspring have already left the nest to overwinter and prepare for the next breeding season.
Natural selection, in this case, would be selecting on those reproductives who are better able to manipulate and control their offspring into helping them. So you have two possible mechanisms that could have driven the development of eusociality — one which confers benefits on an “altruistic” kin group, and one that primarily benefits a “selfish” individual. In the Kapheim et al. study, they decided to “test” these two theories via simulation as well as a manipulated field study of a model organism, a tropical sweat bee called Megalopta genalis. What is unique about this bee is that it can “choose” to be eusocial or not; nest foundresses will remain either solitary (in the case of if their first brood is all male), or they can establish a eusocial colony if their first brood has at least one female. The latter of which, like other eusocial colonies, will be composed primarily of many sterile daughters helping/protecting the foundress and her offspring. Of these cases, about ~6% of eusocial colonies fail, i.e. the daughters, for whatever reason, will all leave to try to reproduce instead of staying to help the mother.
Figure 2: A pinned specimen of the tropical sweat bee (M. genalis).
Because of this “facultative”/optional eusociality, it was assumed the species would make a good model for envisioning an earlier state of eusocial evolution. The researchers therefore carried out a manipulated field study, in which M. genalis foundresses were allowed to establish nests in artificial hives placed throughout their habitat, and whether they became eusocial or remained solitary, their ability to survive to the second brood attempt, genotypes, and other pertinent data were noted and recorded. After calculating fitness costs and benefits for foundresses versus workers based on their observations, they then created an evolutionary simulation, one in which genes were supposed to influence the expression of manipulative behaviour by foundresses and another in which genes were supposed to influence/select for worker/cooperative behaviours. Survivability parameters were set/extrapolated from their field observations (for example, it was assumed approximately ~63% of the starting population would survive to create a first brood, as was found in their field study). This was to model over many generations, given such a situation, whether a eusocial state was more likely to evolve with selection acting on the characteristic of parental parasitism/manipulation, or whether it would be better accounted for by inclusive fitness, with selection acting on worker altruism.
It was found that, given workers are not all as closely-related as 75%, the workers had calculated fitnesses much lower than solitary foundresses, and even lower fitness scores than those reproductives who were mated and tried to establish a nest but still failed (such as by disease). Getting one point for trying is always a leg up on a 0 for absolutely no effort, so to speak. The simulation results were also able to better account for the emergence of eusociality with the selective force reinforcing maternal manipulation, as when workers and their fitness were selected upon, at an equilibrium state “failed eusocial” nests were dominant, which, again, is when the workers would all leave the nest to try to create their own brood. This clearly did not match with the reality, in which such a phenomena is rare (again, only 6% of M. genalis eusocial nests end up failing), and while the worker altruism model could be made to more closely align with observed conditions, it called for more restrictive parameters (such as increasing worker relatedness) which were not required by maternal manipulation. So, while the authors concluded that both inclusive fitness and maternal manipulation likely play some role in the evolution of eusociality — workers aren’t entirely bereft, and the parents clearly stand to gain in their manipulation — it seems likely maternal manipulation/parental parasitism was primarily the driving behaviour early in eusociality’s evolutionary history.
What can be learned about the human condition from any of that? That is for you to discern, but I can speak for what it meant to me — I think it speaks towards, informs about, the human phenomena of “civilisation” and its development. There have been a number of individuals who have, more or less enthusiastically, likened civilisation to eusocial societies amongst insects. There are obviously some important qualitative differences, the nuances that we should never look over entirely in any such comparison — for example, human civilisations obviously aren’t composed of several million brothers and sisters ruled over by a mother and/or father, or several such extremely fecund pairs, and so there is no chance for the proposed prerequisite of strong kin selection via marked genetic relatedness. However, the implication of the referenced study suggests this does not matter much at all, at least in the foundational stages of eusocial evolution: the primary driver is “parental parasitism,” “maternal manipulation,” the “selfish genome” at it again. And you certainly get much of that in human civilisation.
Division of labour, castes divided by their function within the society — just like among the insect counterparts —organised and enforced by social elites, all of it in service to better maximising, via luxurious protection, the elite’s own fitness, regardless of whether or not it bolsters the worker’s own fitness. Rather than using pheromone signals, the method of control and manipulation for humans lies in the unique human attribute of enculturation — the ability to be psychologically shaped to a considerable degree by learning, rather than by lock-and-key, chemical instincts. If you question that the project of such unequal societies is meant to maximise the fitness of only a few, one needs only refer to the fact that almost everyone on the planet is descended in part from historical elites/royals — if you are the maker of the “Law,”if you are the “authority,” it is often only one’s own moral scruples holding one back from mating with (or outright raping) whoever one pleases, while condemning to death — denying them and their genes the ability to carry forth into the future — any and all who displease or threaten you. The genetic record indicates there were not too many in the position who had that sort of self-restraint — and so one finds millions of descendents of Genghis Khan, while John Doe, who died in the trenches of World War 1 as a pawn at age 19, has no progeny to speak of. While it may afford an individual some certain comforts in the short term, in the long term, non-elite lineages have a very high chance of being crumpled up and disposed of, failing, in such a social organisation, whether by being starved to death by the deliberate incompetence or genocidal intentions of one’s superiors, or expected to sacrifice yourself for the good of the “society” (which, again, in such a eusocial state, translates into the good/interests of the elites).
While elites may not actually be your parents, the social behaviours they condition and expect is that of a parent over children. “Civilised” people can readily be likened to children, whatever many philosophical champions of the “civil” have historically liked to claim the inverse, i.e. the “savage” being childlike, embryonic, infantile. The domestic man, much like his subdued cultivars of formerly wild animals, is much more docile, obedient, generally-less intelligent due to giving up much of their executive cognition to the masters of their lives, deferring often or always to the surrogate parent/authority above them (the one who refuses such submission is usually jailed out of existence or executed, as already noted). Through most of human evolution, and very likely amongst our hominid predecessors, the only figure you would really be expected to obey without question was the parent, while peers could be questioned or challenged at your discretion. If the phenomena of eusociality depends on such, i.e. parental manipulation, then the human analogue — civilisation — would require and select for increasing intellectual/social infantilism amongst its subjects, breeding out the more aggressive/less-permissive varieties, in order for the elites to better perfect their parental parasitism.
Figure 3: Representative illustration of the hierarchical structure of ancient Egyptian civilisation. The format of hierarchical/eusocial societies changes little, with the bulk of society forming the base of the metaphorical “pyramid” (slaves or labourers) and a few individuals forming a “privileged” pinnacle. An inverted “wealth” pyramid can be laid over it, in which the majority of the wealth produced by the society belongs to or is at the disposal of the top of the pyramid (and all of the fitness benefits that entails), while the least amount of wealth and consequent lowest fitness is found at the social base.
Are these — docility, obedience, a sacrifice of the executive powers by the individual — not the “virtues” often extolled by many of the forward-thinkers of the modern day, past and present? An abhorrence and denigration of all that is violent, except when it is used — for our own good, supposedly — by the agency of the State, at the discretion of the elites. Praise for the “meek” and the “gentle,” he who accepts his subjugated station whatever the burdens and abuse, with promises of a Kingdom Come (or, more often in the modern day, the flimsier appeals to the good of “Man,” of social “progress”). Boasting over the extraordinary complexity of modern civilisation, in which every man is “freed” from the drudgery of self-sufficiency, of being responsible and executive over much of his own affairs, of which in olden times he would have spent much of his day furbishing the tools or tending the food he would have had to provide for himself; the vast majority of his needs are now built and supplied to him by a huge network of others, his executive, “adult” responsibilities therefore being subsumed into this veritable hive, and for it, he himself can indulge in the liberty of spending most of his waking day working to furbish himself with his basic needs. There may be something to say about diminished intellectual stature there, particularly of those who extol such. And while elites — the State — generally have an incomplete regulation of their workers’ reproduction (again, by being capricious masters of life and death, war and peace), unlike the Hymenopteran analogues, the emergence of various philosophies and movements incorporating eugenic ideals in the past few centuries, as well as the emergence of modern states that have the power to decree limits on reproduction and which can exercise the ability to take away children from parents judged unworthy by State standards, have made it not at all unlikely that such will be the next phase in the human march towards “true sociality” — eusociality. All for the good of the One — all for the fitness of the Elites. O brave new world, that has such people in’t!
There is also something less dire, less pessimistic, to be learned from this look into the Deeps. I would like to draw attention to the “unsuccessful” foundresses having a calculated higher fitness than eusocial workers who forgo their own reproduction. Going your own way — trying, and enduring all the risks of that — is worth more than the fruits of subjugation, of sitting on your hands in comfort, waiting for you or your line’s eventual annihilation for the “good of society.” In the case of the eusocial insects, failing to reproduce cuts them away from the future; they and what they carry within them won’t carry on, that unique spark of life extinguished forever. With eusocial humans, oneself and one’s children might reproduce, and all might seem stable enough, in the scope of short and limited lifespans — but in the scope of evolution, not even a blink of the eye may pass before your descendants may all be living in poverty, freezing to death on the streets, starving in a wartorn country, their spark stamped out, in service to the elites, the ones privileged by their parasitism in being secure in their evolutionary fortunes, at least for a much longer time than those on the bottom of the hierarchy ever will be.
Reclaim your wings and fly forth, making your own bold way, is what I see and learn from the depths of life, in this glance and in many other places I turn my gaze. Strive always to climb to the heights, to reject the role of the inferior or subordinate, and to suggest others of your kith and kin to do the same, for the predators and parasites of life have never shyed away from a free lunch or an easy meal for their own benefit and survival, whether that meal comes in actual flesh or the fruits produced by the work of such flesh. To try and to have dared gets you closer to fortune than to have not dared at all. That is a lesson for all time, for the deeds you accomplish here and now, and for what you leave that shall echo down the evolving ages.
Kapheim, K.M., Nonacs, P., Smith, A.R., Wayne, R.K., and Wcislo, W.T. (2015). Kinship, parental manipulation and evolutionary origins of eusociality. Proceedings of the Royal Society B: Biological Sciences, 282(1803). doi:10.1098/rspb.2014.2886.
Free Full Text: https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4345457/
For an interesting read on the “epidemiological transitions” in human history (major shifts in the patterns of human health and diseases) that includes a brief note on “social parasitism” of elites in human society, I would additionally recommend:
Armelagos, G., Brown, P., and Turner, B. (2005). Evolutionary, Historical and Political Economic Perspectives on Health and Disease. Social Science & Medicine (1982), 61(4), 755-765. doi:10.1016/j.socscimed.2004.08.066.