Darwinian evolution was thought to apply primarily at the individual level. Survival of the fittest implied that selfishness was essential for any organism, and so altruism could not exist. Yet, eusocial organisms, such as ants, wasps, and bees, form hierarchical social systems with reproductive queens and sterile workers. So, many insects are taking the evolutionarily counterintuitive step of sacrificing their own reproduction, and instead caring for the offspring of others. These insects, altruistically contributing to their community’s welfare, not themselves reproducing but sacrificing themselves for the greater good of the colony, are common, but should not exist!
Indeed, eusocial insects are remarkably successful. Just two percent of insects are eusocial, but they consist of over sixty percent of all insect biomass. The biomass of ants alone is more than half that of all insects, exceeding that of all terrestrial nonhuman vertebrates combined. Such selflessness in eusocial insect colonies has been explained by genetic relatedness called kin selection. It was a cornerstone of evolutionary biology, explaining social and cooperative behavior across the animal kingdom, even in humans. The trouble with kin selection is that it simply does not fit the data, according to Harvard scientists:
For the past four decades kin selection theory… has been the major theoretical attempt to explain the evolution of eusociality. Here we show the limitations of its approach.Prof E O Wilson
When eusociality happens in insect species that have a colony of siblings, all with the same mother, the queen, they are plainly closely related, having a majority of the same genes. But eusociality in insect species in which siblings were less related to each other, and even in a mammal, the naked mole rat, showed that high genetic relatedness, “inclusive fitness”, and eusociality no longer need correlate. Wilson, the Pellegrino University Professor, Emeritus, at Harvard says:
The empirical evidence gathered in our paper demonstrates that eusociality is exceedingly rare because species must navigate a lengthy “evolutionary labyrinth” to reach this state. We hope our new theory for the evolution of eusociality will open up sociobiology to new avenues of research by liberating the study of social evolution from mandatory adherence to kin selection theory. After four decades ruling the roost, it is time to recognize this theory’s very limited prowess.
The new theory proposed by evolutionary biologist, Wilson, a founder of modern sociobiology, and mathematical biologists, Martin A Nowak and Corina E Tarnita, shows that straightforward natural selection theory alone can explain the evolution of eusocial behavior, without the need for kin selection theory.
Nowak, professor of mathematics and of biology at Harvard and director of the university’s Program for Evolutionary Dynamics says that inclusive fitness theory (IFT), which tries to calculate fitness effects conferred on relatives, sometimes may be a suitable alternative to standard population genetics, but generally is not. This analysis shows that IFT rests on assumptions which rarely hold in Nature. Contrary to many previous claims, IFT is not an extended theory of evolution and is not needed to explain eusociality. Standard natural selection theory represents a simpler and superior approach, and provides an exact framework for interpreting empirical observations. Tarnita, a junior fellow in Harvard’s Society of Fellows says:
Inclusive fitness theory is almost like a shortcut. It only applies to a small subset of all possible models. Outside of that subset, it doesn’t work.
Eusociality is rare, but important in evolutionary biology because the few species that adhere to it—including social insects and, to an extent, humans—rank among the planet’s most dominant. Humans, who are loosely eusocial, dominate land vertebrates. Tarnita explained:
Eusociality has arisen independently some 10 to 20 times in the course of evolution. Our model shows that it is difficult to get eusociality in the first place, but that it is very stable once it is established. A colony behaves like a “superorganism”, reproducing the genome of the queen and the sperm she has stored.
From this perspective, a worker ant is something like a cell—part of a larger evolutionary unit, not a unit unto itself:
Our model proves that looking at a worker ant and asking why it is altruistic is the wrong level of analysis. The important unit is the colony.
So, the researchers offered their alternative theory. A primordial, solitary ant, that lived near a food source—something like the primitive Martialis heureka—developed genetic mutations causing it to feed its offspring, rather than leaving them fend for themselves. This “progressive provisioning” is widespread in insects. From this single founder, selection moves to the level of colony. Another mutation kept offspring near the nest rather than leaving. The insects have natural instincts, even as solitary animals, to do certain things. Nowak continued:
Put two normally solitary wasps together, and if one builds a hole, the other puts an egg in it.
The other sees the egg, and feeds it. This alone is enough to start a small colony, from which eusociality could emerge from further mutations bringing specialization, reproduction restricted to queens alone, and favoring the colony’s success. Within this colony, a queen would be analogous to a human egg or sperm cell—a unit that embodies the whole. Worker self sacrifice in such a unit is no more nonsensical than that of a white blood cell.
In summary, the proposal has three distinct steps species can take to sidestep eusociality’s evolutionary cost:
- species must form groups within a population, as when nests or food attract individuals to discrete locations some distance apart, when parents and offspring remain together, or when migrating flocks follow leaders
- species must accumulate traits, arising through ordinary natural selection, that favor the switch to eusociality, as when Ceratina and Lasioglossum bees, on the cusp of eusociality, cooperate in foraging, tunneling, and guarding resources, and progressive provisioning, in which a female builds a nest, lays an egg in it, and then feeds or guards larvae until they mature. The candidate species must build a defensible nest
- individuals must develop genes supporting eusociality, whether by mutation or recombination. Crossing the threshold to eusociality essentially requires that a female and her offspring not disperse to start new, individual nests, but rather remain at the old nest. While eusocial genes have yet to be identified, at least two eusocial ant species are known to have genes that quell the urge to roam from the nest.
If these steps lead to a species becoming eusocial, the evolutionary costs of individuals foregoing reproduction are compensated by the reduced mortality of the queen and her larvae, which are protected by the colony. In some ant species, a queen, that alone might live for only a few months, can live for over 25 years as part of a colony, producing millions of offspring in the process.
The new theory explains everything that kin selection does, plus what it does not. The set of steps is the evolutionary “labyrinth”, the navigation of which is difficult and so rarely happens. So, eusociality is rare. But neither kin selection nor inclusive fitness need be invoked, either in eusociality, or in any cooperative behavior. Tarnita said the new theory of eusociality may help describe how single celled organisms gave rise to multicellular organisms.
Approving the idea, David Sloan Wilson, an evolutionary biologist at Binghamton University, said everything nice about human behavior—based on interactions among genetic relatives during the Stone Age, and now being incorrectly expressed—is pathetic as an explanatory framework. Especially troubling to him is that kin selection, with its intuitive appeal to our preference for family over strangers, has been applied to human social life. Tarnita comments that human selflessness and cooperation is of a different sort, involving the interaction of culture and sentience, not just genetics and environment:
There are certain things we can learn from ants, but I wouldn’t try to draw a parallel. It’s easier to think about ants, but people are complicated.
Disapproving, Rice University’s David Queller thought the new model “involves, and I suspect requires, close kinship”.
The reaction is mixed, and there will doubtless be volleys from either side to come. It seems likely, though, that the original small colony would arise from a family grouping, so the participants will be kin, even if the model could notionally work with unrelated solitary insects. The authors do not totally write off kin selection, simply considering it to apply to a “subset” of models of eusocial evolution. Maybe the subset is necessary, if not sufficient, for any species to enter the “labyrinth”!