Group selection

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Early explanations of social behavior, such as the lekking of blackcock, spoke of "the good of the species".^[1] Blackcocks at the Lek watercolour and bodycolour by Archibald Thorburn, 1901.

Group selection is a proposed mechanism of evolution in which natural selection acts at the level of the group, instead of at the more conventional level of the individual.

Early authors such as V. C. Wynne-Edwards and Konrad Lorenz argued that the behavior of animals could affect their survival and reproduction as groups, speaking for instance of actions for the good of the species. From the mid 1960s, evolutionary biologists such as John Maynard Smith argued that natural selection acted primarily at the level of the individual. They argued on the basis of mathematical models that individuals would not altruistically sacrifice fitness for the sake of a group. They persuaded the majority of biologists that group selection did not occur, other than in special situations such as the haplodiploid social insects like honeybees (in the Hymenoptera), where kin selection was possible.

In 1994 David Sloan Wilson and Elliott Sober argued for multi-level selection, including group selection, on the grounds that groups, like individuals, could compete. In 2010 three authors including E. O. Wilson, known for his work on social insects especially ants, again revisited the arguments for group selection. They argued that group selection can occur when competition between two or more groups, some containing altruistic individuals who act cooperatively together, is more important for survival than competition between individuals within each group. Their proposals provoked a strong rebuttal from a large group of evolutionary biologists.^[2]

As of yet, there is no clear consensus among biologists regarding the importance of group selection. Steven Pinker expressed his ambivalence with the theory: "Human beings live in groups, are affected by the fortunes of their groups, and sometimes make sacrifices that benefit their groups. Does this mean that the human brain has been shaped by natural selection to promote the welfare of the group in competition with other groups, even when it damages the welfare of the person and his or her kin?... I think that this reasonableness is an illusion. The more carefully you think about group selection, the less sense it makes, and the more poorly it fits the facts of human psychology and history."^[3] However, there is active debate among specialists in many fields of study. It is possible that a theory of group selection can be modified to provide valuable explanations. Group selection could be useful for understanding the evolution of human culture, since humans form groups that are unlike any other animal. Group selection may be used to understand human history. Some researchers have used the framework to understand the development of human morality.

Early Developments

Charles Darwin developed the theory of evolution in his book, Origin of Species. Darwin also made the first suggestion of group selection in The Descent of Man that the evolution of groups could affect the survival of individuals. He wrote, "If one man in a tribe... invented a new snare or weapon, the tribe would increase in number, spread, and supplant other tribes. In a tribe thus rendered more numerous there would always be a rather better chance of the birth of other superior and inventive members."^[4][5]

Once Darwinism had been accepted, animal behavior was glibly explained with unsubstantiated hypotheses about survival value, which was largely taken for granted. The naturalist Konrad Lorenz had argued loosely in books like On Aggression (1966) that animal behavior patterns were "for the good of the species",^[1][6] without actually studying survival value in the field;^[6] Richard Dawkins noted that Lorenz was a "'good of the species' man"^[7] so accustomed to group selection thinking that he did not realize his views "contravened orthodox Darwinian theory".^[7] The ethologist Niko Tinbergen praised Lorenz for his interest in the survival value of behavior, and naturalists enjoyed Lorenz's writings for the same reason.^[6] In 1962, group selection was used as a popular explanation for adaptation by the zoologist V. C. Wynne-Edwards.^[8][9] In 1976, Richard Dawkins wrote a well-known book on the importance of evolution at the level of the gene or the individual, The Selfish Gene.^[10]
 

Social behavior in honeybees is explained by kin selection: their haplodiploid inheritance system makes workers very closely related to their queen (centre).

From the mid 1960s, evolutionary biologists argued that natural selection acted primarily at the level of the individual. In 1964, John Maynard Smith,^[11] C. M. Perrins (1964),^[12] and George C. Williams in his 1966 book Adaptation and Natural Selection cast serious doubt on group selection as a major mechanism of evolution; Williams's 1971 book Group Selection assembled writings from many authors on the same theme.^[13][14]

It was at that time generally agreed that the primary exception of social group selection was in the social insects, and the explanation was limited to the unique inheritance system (involving haplodiploidy) of the eusocial Hymenoptera such as honeybees, which encourages kin selection, since workers are closely related.^[2]

The great majority of evolutionary biologists believe (2011) that selection above the level of the individual is a special case, probably limited to the unique inheritance system (involving haplodiploidy) of the eusocial Hymenoptera such as honeybees, which may encourage kin selection since workers are closely related.^[2]

Kin selection and inclusive fitness theory

Early group selection models assumed that genes acted independently, for example a gene that coded for cooperation or altruism. Genetically-based reproduction of individuals implies that, in group formation, the altruistic genes would need a way to act for the benefit of members in the group to enhance the fitness of many individuals with the same gene.^[15] But it is expected from this model that individuals of the same species would compete against each other for the same resources. This would put cooperating individuals at a disadvantage, making genes for cooperation tend to be eliminated. Group selection on the level of the species is flawed because it is difficult to see how selective pressures would be applied to competing/non-cooperating individuals.

Experiments from the late 1970s suggested that selection involving groups was possible.^[16] Kin selection between related individuals is accepted as an explanation of altruistic behavior. In this model, genetically related individuals cooperate because survival advantages to one individual also benefit kin who share some fraction of the same genes, giving a mechanism for favoring genetic selection.^[17]

Inclusive fitness theory, first proposed by W. D. Hamilton in the early 1960s, gives a selection criterion for evolution of social traits when social behavior is costly to an individual organism's survival and reproduction. This behavior could emerge under conditions such that the statistical likelihood that benefits accrue to the survival and reproduction of other organisms whom also carry the social trait. Inclusive fitness theory is a general treatment of the statistical probabilities of social traits accruing to any other organisms likely to propagate a copy of the same social trait. Kin selection theory treats the narrower but simpler case of the benefits to close genetic relatives (or what biologists call 'kin') who may also carry and propagate the trait. A significant group of biologists support inclusive fitness as the explanation for social behavior in a wide range of species, as supported by experimental data. An article was published in Nature with over a hundred coauthors.^[2]

One of the questions about kin selection is the requirement that individuals must know if other individuals are related to them, or kin recognition. Any altruistic act has to preserve similar genes. One argument given by Hamilton is that many individuals operate in "viscous" conditions, so that they live in physical proximity to relatives. Under these conditions, they can act altruistically to any other individual, and it is likely that the other individual will be related. This population structure builds a continuum between individual selection, kin selection, kin group selection and group selection without a clear boundary for each level. However, early theoretical models by D.S. Wilson et al.^[18] and Taylor^[19] showed that pure population viscosity cannot lead to cooperation and altruism. This is because any benefit generated by kin cooperation is exactly cancelled out by kin competition; additional offspring from cooperation are eliminated by local competition. Mitteldorf and D. S. Wilson later showed that if the population is allowed to fluctuate, then local populations can temporarily store the benefit of local cooperation and promote the evolution of cooperation and altruism.^[20] By assuming individual differences in adaptations, Yang further showed that the benefit of local altruism can be stored in the form of offspring quality and thus promote the evolution of altruism even if the population does not fluctuate. This is because local competition among more individuals resulting from local altruism increases the average local fitness of the individuals that survive.^[21]

Another explanation for the recognition of genes for altruism is that a single trait, group reciprocal kindness, is capable of explaining the vast majority of altruism that is generally accepted as "good" by modern societies. The phenotype of altruism relies on recognition of the altruistic behavior by itself. The trait of kindness will be recognized by sufficiently intelligent and undeceived organisms in other individuals with the same trait. Moreover, the existence of such a trait predicts a tendency for kindness to unrelated organisms that are apparently kind, even if the organisms are of a completely different species. The gene need not be exactly the same, so long as the effect or phenotype is similar. Multiple versions of the gene—or even meme—would have virtually the same effect. This explanation was given by Richard Dawkins as an analogy of a man with a green beard. Green-bearded men tend to cooperate with each other simply by seeing a green beard, where the green beard trait is incidentally linked to the reciprocal kindness trait.^[10]

Multilevel selection theory

Kin selection or inclusive fitness is accepted as an explanation for cooperative behavior in many species, but there are some species, including some human behavior, that are difficult to explain with only this approach. In particular, it doesn't seem to explain the cause of the (relatively) rapid rise of human civilization. David Sloan Wilson has argued that other factors must also be considered in evolution.^[22] Since the 1990s, group selection models have seen a resurgence and further refinement.


Early group selection models were flawed because they assumed that genes acted independently; but genetically-based interactions among individuals are ubiquitous in group formation because genes must cooperate for the benefit of association in groups to enhance the fitness of group members.^[28] Additionally, group selection on the level of the species is flawed because it is difficult to see how selective pressures would be applied; selection in social species of groups against other groups, rather than the species entire, seems to be the level at which selective pressures are plausible. On the other hand, kin selection is accepted as an explanation of altruistic behavior.^[17][29] Some biologists argue that kin selection and multilevel selection are both needed to "obtain a complete understanding of the evolution of a social behavior system".^[30]


In 1994 David Sloan Wilson and Elliott Sober argued that the case against group selection had been overstated. They considered whether groups can have functional organization in the same way as individuals, and consequently whether groups can be "vehicles" for selection. They do not posit evolution on the level of the species, but selective pressures that winnow out small groups within a species, e.g. groups of social insects or primates. Groups that cooperate better might survive and reproduce more than those that did not. Resurrected in this way, Wilson & Sober's new group selection is called multilevel selection theory.^[31]

In 2010, M. A. Nowak, C. E. Tarnita and E. O. Wilson argued for multi-level selection, including group selection, to correct what they saw as deficits in the explanatory power of inclusive fitness.^[32] The response was a back-lash from 137 other evolutionary biologists who argued "that their arguments are based upon a misunderstanding of evolutionary theory and a misrepresentation of the empirical literature".^[2]
 

David Sloan Wilson and Elliott Sober's 1994 Multilevel Selection Model, illustrated by a nested set of Russian matryoshka dolls. Wilson himself compared his model to such a set.

Wilson compared the layers of competition and evolution to nested sets of Russian matryoshka dolls.^[23] The lowest level is the genes, next come the cells, then the organism level and finally the groups. The different levels function cohesively to maximize fitness, or reproductive success. The theory asserts that selection for the group level, involving competition between groups, must outweigh the individual level, involving individuals competing within a group, for a group-benefiting trait to spread.^[33]

Multilevel selection theory focuses on the phenotype because it looks at the levels that selection directly acts upon.^[23] For humans, social norms can be argued to reduce individual level variation and competition, thus shifting selection to the group level. The assumption is that variation between different groups is larger than variation within groups. Competition and selection can operate at all levels regardless of scale. Wilson wrote, "At all scales, there must be mechanisms that coordinate the right kinds of action and prevent disruptive forms of self-serving behavior at lower levels of social organization."^[22] E. O. Wilson summarized, "In a group, selfish individuals beat altruistic individuals. But, groups of altruistic individuals beat groups of selfish individuals."

Wilson ties the multilevel selection theory regarding humans to another theory, gene-culture coevolution, by acknowledging that culture seems to characterize a group-level mechanism for human groups to adapt to environmental changes.^[33]

MLS theory can be used to evaluate the balance between group selection and individual selection in specific cases.^[33] An experiment by William Muir compared egg productivity in hens, showing that a hyper-aggressive strain had been produced through individual selection, leading to many fatal attacks after only six generations; by implication, it could be argued that group selection must have been acting to prevent this in real life.^[36] Group selection has most often been postulated in humans and, notably, eusocial Hymenoptera that make cooperation a driving force of their adaptations over time and have a unique system of inheritance involving haplodiploidy that allows the colony to function as an individual while only the queen reproduces.^[37]

Wilson and Sober's work revived interest in multilevel selection. In a 2005 article,^[38] E. O. Wilson argued that kin selection could no longer be thought of as underlying the evolution of extreme sociality, for two reasons. First, he suggested, the argument that haplodiploid inheritance (as in the Hymenoptera) creates a strong selection pressure towards nonreproductive castes is mathematically flawed.^[39] Second, eusociality no longer seems to be confined to the hymenopterans; increasing numbers of highly social taxa have been found in the years since Wilson's foundational text Sociobiology: A New Synthesis was published in 1975.^[40] These including a variety of insect species, as well as two rodent species (the naked mole-rat and the Damaraland mole rat). Wilson suggests the equation for Hamilton's rule:^[41]

rb > c

(where b represents the benefit to the recipient of altruism, c the cost to the altruist, and r their degree of relatedness) should be replaced by the more general equation

rb_k + b_e > c

in which b_k is the benefit to kin (b in the original equation) and b_e is the benefit accruing to the group as a whole. He then argues that, in the present state of the evidence in relation to social insects, it appears that b_e>rb_k, so that altruism needs to be explained in terms of selection at the colony level rather than at the kin level. However, kin selection and group selection are not distinct processes, and the effects of multi-level selection are already accounted for in Hamilton's rule, rb>c,^[42] provided that an expanded definition of r, not requiring Hamilton's original assumption of direct genealogical relatedness, is used, as proposed by E. O. Wilson himself.^[43]

Spatial populations of predators and prey show restraint of reproduction at equilibrium, both individually^[44] and through social communication,^[45] as originally proposed by Wynne-Edwards. While these spatial populations do not have well-defined groups for group selection, the local spatial interactions of organisms in transient groups are sufficient to lead to a kind of multi-level selection. There is however as yet no evidence that these processes operate in the situations where Wynne-Edwards posited them.^{[citation needed]}

Rauch et al.'s analysis^[44] of a host-parasite situation, which was recognised as one where group selection was possible even by E. O. Wilson (1975), is broadly hostile to the whole idea of group selection.^[40] Specifically, the parasites do not individually moderate their transmission; rather, more transmissible variants "continually arise and grow rapidly for many generations but eventually go extinct before dominating the system."

Applications

Differing evolutionarily stable strategies

The problem with group selection is that for a whole group to get a single trait, it must spread through the whole group first by regular evolution. But, as J. L. Mackie suggested, when there are many different groups, each with a different evolutionarily stable strategy, there is selection between the different strategies, since some are worse than others.^[46] For example, a group where altruism was universal would indeed outcompete a group where every creature acted in its own interest, so group selection might seem feasible; but a mixed group of altruists and non-altruists would be vulnerable to cheating by non-altruists within the group, so group selection would collapse.^[47]

Implications in population biology

Social behaviors such as altruism and group relationships can impact many aspects of population dynamics, such as intraspecific competition and interspecific interactions. In 1871, Darwin argued that group selection occurs when the benefits of cooperation or altruism between subpopulations are greater than the individual benefits of egotism within a subpopulation.^[48] This supports the idea of multilevel selection, but kinship also plays an integral role because many subpopulations are composed of closely related individuals. An example of this can be found in lions, which are simultaneously cooperative and territorial.^[49] Within a pride, males protect the pride from outside males, and females, who are commonly sisters, communally raise cubs and hunt. However, this cooperation seems to be density dependent. When resources are limited, group selection favors prides that work together to hunt. When prey is abundant, cooperation is no longer beneficial enough to outweigh the disadvantages of altruism, and hunting is no longer cooperative.^[49]

Interactions between different species can also be affected by multilevel selection. Predator-prey relationships can also be affected. Individuals of certain monkey species howl to warn the group of the approach of a predator.^[50] The evolution of this trait benefits the group by providing protection, but could be disadvantageous to the individual if the howling draws the predator's attention to them. By affecting these interspecific interactions, multilevel and kinship selection can change the population dynamics of an ecosystem.^[50]

Multilevel selection attempts to explain the evolution of altruistic behavior in terms of quantitative genetics. Increased frequency or fixation of altruistic alleles can be accomplished through kin selection, in which individuals engage in altruistic behavior to promote the fitness of genetically similar individuals such as siblings. However, this can lead to inbreeding depression,^[51] which typically lowers the overall fitness of a population. However, if altruism were to be selected for through an emphasis on benefit to the group as opposed to relatedness and benefit to kin, both the altruistic trait and genetic diversity could be preserved. However, relatedness should still remain a key consideration in studies of multilevel selection. Experimentally imposed multilevel selection on Japanese quail was more effective by an order of magnitude on closely related kin groups than on randomized groups of individuals.^[52]

Gene-culture coevolution in humans

Humanity has developed extremely rapidly, arguably through gene-culture coevolution, leading to complex cultural artefacts like the gopuram of the Sri Mariammam temple, Singapore.

Gene-culture coevolution (also called dual inheritance theory) is a modern hypothesis (applicable mostly to humans) that combines evolutionary biology and modern sociobiology to indicate group selection.^[53] It treats culture as a separate evolutionary system that acts in parallel to the usual genetic evolution to transform human traits.^[54] It is believed that this approach of combining genetic influence with cultural influence over several generations is not present in the other hypotheses such as reciprocal altruism and kin selection, making gene-culture evolution one of the strongest realistic hypotheses for group selection. Fehr provides evidence of group selection taking place in humans presently with experimentation through logic games such as prisoner’s dilemma, the type of thinking that humans have developed many generations ago.^[55]

Gene-culture coevolution allows humans to develop highly distinct adaptations to the local pressures and environments more quickly than with genetic evolution alone. Robert Boyd and Peter J. Richerson, two strong proponents of cultural evolution, postulate that the act of social learning, or learning in a group as done in group selection, allows human populations to accrue information over many generations.^[56] This leads to cultural evolution of behaviors and technology alongside genetic evolution. Boyd and Richerson believe that the ability to collaborate evolved during the Middle Pleistocene, a million years ago, in response to a rapidly changing climate.^[56]

In 2003, Herbert Gintis examined cultural evolution statistically, offering evidence that societies that promote pro-social norms have higher survival rates than societies that do not.^[57]

Gintis wrote that genetic and cultural evolution can work together. Genes transfer information in DNA, and cultures transfer information encoded in brains, artifacts, or documents. Language, tools, lethal weapons, fire, cooking, etc., have a long-term effect on genetics. For example, cooking led to a reduction of size of the human gut, since less digestion is needed for cooked food. Language led to a change in the human larynx and an increase in brain size. Projectile weapons led to changes in human hands and shoulders, such that humans are much better at throwing objects than the closest human relative, the chimpanzee.^[58]

Criticism

The use of the Price equation to support group selection was challenged by van Veelen in 2012, arguing that it is based on invalid mathematical assumptions.^[59]

Richard Dawkins and other advocates of the gene-centered view of evolution remain unconvinced about group selection.^[60][61][62] In particular, Dawkins suggests that group selection fails to make an appropriate distinction between replicators and vehicles.^[63]

The psychologist Steven Pinker concluded that "group selection has no useful role to play in psychology or social science", as it "is not a precise implementation of the theory of natural selection, as it is, say, in genetic algorithms or artificial life simulations. Instead it is a loose metaphor, more like the struggle among kinds of tires or telephones."^[3]

The evolutionary biologist Jerry Coyne summarized the arguments in The New York Times in non-technical terms as follows:^[64]

Group selection isn't widely accepted by evolutionists for several reasons. First, it's not an efficient way to select for traits, like altruistic behavior, that are supposed to be detrimental to the individual but good for the group. Groups divide to form other groups much less often than organisms reproduce to form other organisms, so group selection for altruism would be unlikely to override the tendency of each group to quickly lose its altruists through natural selection favoring cheaters. Further, little evidence exists that selection on groups has promoted the evolution of any trait. Finally, other, more plausible evolutionary forces, like direct selection on individuals for reciprocal support, could have made humans prosocial. These reasons explain why only a few biologists, like [David Sloan] Wilson and E. O. Wilson (no relation), advocate group selection as the evolutionary source of cooperation.

Are We Enlightened Guardians, Or Are We Apes Designing Humans?

May 22, 2006 by Douglas Mulhall
Original link:  http://www.kurzweilai.net/are-we-enlightened-guardians-or-are-we-apes-designing-humans
Originally published in Nanotechnology Perceptions: A Review of Ultraprecision Engineering and Nanotechnology, Volume 2, No. 2, May 8, 2006. Reprinted with permission on KurzweilAI.net, May 22, 2006.

Thanks in part to molecular manufacturing, accelerated developments in AI and brain reverse-engineering could lead to the emergence of superintelligence in just 18 years. Are we ready for the implications — like possible annihilation of Homo sapiens? And will we seem to superintelligence what our ape-like ancestors seem to us: primitive?

Most students of artificial intelligence are familiar with this forecast made by Vernor Vinge in 1993¹: "Within thirty years, we will have the technological means to create superhuman intelligence. Shortly after, the human era will be ended."

That was thirteen years ago. Many proponents of super-intelligence say we are on track for that deadline, due to the rate of computing and software advances. Skeptics argue this is nonsense and that we’re still decades away from it.

But fewer and fewer argue that it won’t happen by the end of this century. This is because history has shown the acceleration of technology to be exponential, as explained in well-known works by inventors such as Ray Kurzweil and Hans Moravec, some of which are elucidated in this volume of essays.

A classic example of technology acceleration is the mapping of the human genome, which achieved most of its progress in the late stages of a multi-year project that critics wrongly predicted would take decades. The rate of mapping at the end of the project was exponential compared to the beginning, due to rapid automation that has since transformed the biotechnology industry.

The same may be true of molecular manufacturing (MM) as self-taught machines learn via algorithms to do things faster, better, and cheaper. I won’t describe the technology of MM here because that is well covered in other essays by more competent experts.

MM is important to super-intelligence because it will revolutionize the processes required to understand our own intelligence, such as neural mapping via neural probes that non-destructively map the brain. It also will accelerate three-dimensional computing, where the space between computing units is reduced and efficiency multiplied in the same way that our own brains have done it. Once this happens, the ability to mimic the human brain will accelerate, and self-aware intelligence may follow quickly.

This type of acceleration suggests that Vinge’s countdown to the beginning of the end of the human era must be taken seriously.

The pathways by which super-human intelligence could evolve have been well explained by others and include: computer-based artificial intelligence, bioelectronic AI that develops super-intelligence on its own, or human intelligence that is accelerated or merged with AI. Such intelligence might be an enhancement of Homo sapiens, i.e. part of us, or completely separate from us, or both.

Many experts argue that each of these forms of super-intelligence will enhance humans, not replace them, and although they might seem alien to unenhanced humans, they will still be an extension of us because we are the ones who designed them.

The thought behind this is that we will go on as a species.

Critics, however, point to a fly in that ointment. If the acceleration of computing and software continues apace, then super-intelligence, once it emerges, could outpace Homo sapiens, with or without piggybacking on human intelligence.

This would see the emergence of a new species, perhaps similar in some ways, but in other ways fundamentally different from Homo sapiens in terms of intelligence, genetics, and immunology.

If that happens, the gap between Homo sapiens and super-intelligence could quickly become as wide as the gap between apes and Homo sapiens.

Optimists say this won’t happen, because everybody will get an upgrade simultaneously when super-intelligence breaks out.

Pessimists say that just a few humans or computers will acquire such intelligence first, and then use it to subjugate the rest of us Homo sapiens.

For clues as to who might be right, let’s look at outstanding historical examples of how we’ve used technology and our own immunology in relation to less technologically adept societies, and in relation to other species.

When technologically superior Europeans arrived in North and South America, the indigenous populations didn’t have much time to contemplate such implications because in a just few years, most who came in contact with Europeans were dead from disease. Many who died never laid eyes on a European, as death spread so quickly ahead of the conquerors through unknowing victims.

Europeans at first had no idea that their own immunity to disease would give them such an advantage, but when they realized it, they did everything to use it as a weapon. They did the same with technologies that they consciously invented and knew were superior.

The rapid death of these ancient civilizations, numbering in the tens of millions of persons across two continents, is not etched into the consciousness of contemporary society because those cultures left few written records and had scant time to document their own demise. Most of what they put to pictures or symbols was destroyed by religious zealots or wealth-seeking exploiters.

And so, these civilizations passed quietly into history, leaving only remnants.

By inference, enhanced intelligence easily could take choices about our future out of our hands, and may also be immune to hazards such as mutating viruses that pose dire threats to human society.

Annihilation of Homo sapiens could occur in one of many ways:

• The "oops" factor: accidental annihilation at the hands of a very smart klutz, e.g. by something that is unwittingly immune to things that kill us, or that is smart in one way, but inept in others. Predecessors to super-intelligence may only be smarter than us in some ways, and therein lies a danger. An autistic intelligence could do us in by accident. Just look at current technology, where computers are more capable than humans in some ways but hopeless in others.
• Annihilation in the crossfire of a war-like competition between competing forms of super-intelligence, some of which might include upgraded Homo sapiens. One of the early, deadlier competitions could be for resources as various forms of super-intelligence gobble up space that we occupy, or remake our ecology into an environment more suitable to their needs.
• Deliberate annihilation or assimilation because we are deemed inferior.

If Vernor Vinge is right, we have 18 years before we will face such realities. Centuries ago, the fate of Indian civilizations in North and South America was decided in a similar time span. So, the time to address such risks is now.

This is especially true because paradigms shift more quickly now; therefore, when the event occurs we’ll have less time, perhaps five years or even just one, to consider our options.

What might we use as protection against these multi-factorial threats?

Sun Microsystems’ cofounder Bill Joy’s April 2000 treatise, "Why the future doesn’t need us,"² summarized one field of thought, arguing the case for relinquishment– eschewing certain technologies due to their inherent risks.

Since that time, most technology proponents have been arguing why relinquishment is impractical. They contend that the march of technology is relentless and we might as well go along for the ride, but with safeguards built in to make sure things don’t get too crazy.

Nonetheless, just how we build safeguards into something smarter than us, including an upgraded version of ourselves, has as yet gone unanswered. To see where the solutions might lie, let’s again look at the historical perspective.

If we evaluate the arguments between technology optimists and relinquishment pessimists in relation to the history of the natural world, it becomes apparent that we are stuck between a rock and a hard place.

The ‘rock’ in this case could be an asteroid or comet. If we were to relinquish our powerful new technologies, chances are good that an asteroid would eventually collide with Earth, as has occurred before, thus throwing human civilization back to the dark ages or worse.

For those who scoff at this as an astronomical long shot, be reminded that Comet Shoemaker-Levy 9 punched Earth-sized holes in Jupiter less than a decade after the space tools necessary to witness such events were launched, and just when most experts were forecasting such occurrences to be once-in-a-million-year events that we would likely never see.

Or perhaps we would be thrown back by other catastrophic events that have occurred historically, such as naturally induced climate changes triggered by super-volcanos, collapse of the magnetosphere, or an all-encompassing super-nova.

Due to those natural risks, I argue in my book, Our Molecular Future, that we may have no choice but to proceed with technologies that could just as easily destroy us as protect us.

Unfortunately, as explained in the same book, an equally bad "hard place" sits opposite the onrushing "rock" that threatens us. The hard place is our social ineptness.

In the 21st century, despite tremendous progress, we still do amazingly stupid things. We prepare poorly for known threats including hurricanes and tsunamis. We go to war over outdated energy sources such as oil, and some of us increasingly overfeed ourselves while hundreds of millions of people ironically starve. We often value conspicuous consumption over saving impoverished human lives, as low income victims of AIDS or malaria know too well.

Techno-optimists use compelling evidence to argue that we are vanquishing these shortcomings and that new technologies will overcome them completely. But one historical trend bodes against this: emergence of advanced technologies has been overwhelmingly bad for many of the less intelligent species on Earth.

To cite a familiar refrain: We are massacring millions of wild animals and destroying their habitat. We keep billions more domestic farm animals under inhumane, painful, plague-breeding conditions in increasingly vast numbers.

The depth and breadth of this suffering is so vast that we often ignore it, perhaps because it is too terrible to contemplate. When it gets too bothersome, we dismiss it as animal rights extremism. Some of us rationalize it by arguing that nature has always extinguished species, so we are only fulfilling that natural role.

But at its core lies a searing truth: our behavior as guardians of less intelligent species, which we know feel pain and suffering, has been and continues to be atrocious.

If this is our attitude toward less intelligent species, why would the attitude of superior intelligence toward us be different? It would be foolish to assume that a more advanced intelligence than our own, whether advanced in all or in only some ways, will behave benevolently toward us once it sees how we treat other species.

We therefore must consider that a real near-term risk to our civilization is that we invent something which looks at our ways of treating less intelligent species and decides we’re not worth keeping, or if we are worth keeping, we should be placed in zoos in small numbers where we can’t do more harm. Resulting questions:

• How do we instill into super-intelligence ‘ethical’ behavior that we ourselves poorly exhibit?
• How do we make sure that super-intelligence rejects certain unsavory practices as we banned slavery?
• Can we reach into the future to prevent a super-intelligence from changing its mind about those ethics?

These questions have been debated, but no broad-based consensus has emerged. Instead, as the discussions run increasingly in circles, they suggest that we as a species might be comparable to ‘apes designing humans’.

The ape-like ancestors of Homo sapiens had no idea they were contributing DNA to a more intelligent species. Nor could they hope to comprehend it. Likewise, can we Homo sapiens expect to comprehend what we are contributing to a super-intelligent species that follows us?

As long as we continue to exercise callous neglect as guardians of species less intelligent than ourselves, it could be argued that we are much like our pre-human ancestors: incapable of consciously influencing what comes after us.

The guardianship issue leads to another question: How well are we balancing technology advantages against risks?

In the mere 60 years since our most powerful weapons—nuclear bombs—were invented, we’ve kept them mostly under wraps and congratulated ourselves for that, but we have also seen them proliferate from at first just one country to at least ten, with some of those balanced on the edge of chaos.

Likewise, in the nanoscale technology world that precedes molecular manufacturing, we’ve begun assessing risks posed to human health by engineered nanoparticles, but those particles are already being put into our environment and into us.

In other words, we are still closing the proverbial barn doors after the animals have escaped. This limited level of foresight is light years away from being able to assess how to control the onrushing risks of molecular manufacturing or of enhanced intelligence.

Many accomplished experts have pointed out that the same empowerment of individuals by technologies such as the Internet and biotech could make unprecedented weapons available to small disaffected groups.

Technology optimists argue that this has occurred often in history: new technologies bring new pros and cons, and after we make some awful mistakes with them, things get sorted out.

However, in this case the acceleration rate by its nature puts these technologies in a class of their own, because the evidence suggests they are running ahead of our capacities to contain or balance them. Moreover, the number of violently disaffected groups in our society who could use them is substantial.

To control this, do we need a “pre-crime” capacity as envisaged in the film Minority Report, where Big Brother methods are applied to anticipate crime and strike it down preemptively?

The pros and cons of preemptive strikes have been well elucidated recently. The idea of giving up our freedom in order to preserve our freedom from attack by disaffected groups is being heavily debated right now, without much agreement.

However, one thing seems to have been under-emphasized in these security debates:

Until we do the blatantly positive things such as eliminate widespread diseases, feed the starving, house the homeless, disenfranchise dictators, stop torture, stop inhumane treatment of less intelligent species, and other do-good things that are treated today like platitudes, we will not get rid of violently disaffected groups.

By doing things that are blatantly humane, (despite the efforts of despots and their extremist anti-terrorist counterparts to belittle them as wimpy) we might accomplish two things at once: greatly reduce the numbers of violently disaffected groups, and present ourselves to super-intelligence as being enlightened guardians.

Otherwise, if we continue along the present path, we may someday seem to superintelligence what our ape-like ancestors seem to us: primitive.

In deciding what to do about Homo sapiens, a superior form of intelligence might first evaluate our record as guardians, such as how we treat species less intelligent than ourselves, and how we treat members of our same species that are less technologically adept or just less fortunate.

Why might super-intelligences look at this first? Because just as we are guardians of those less intelligent or fortunate than us, so super-intelligences will be the guardians of us and of other less intelligent species. Super-intelligences will have to decide what to do with us, and with them.

If Vinge is accurate in his forecast, we don’t have much time to set these things straight before someone or something superior to us makes a harsh evaluation.

Being nice to dumb animals or poor people is by no means the only way of assuring survival of our species in the face of something more intelligent than us. Using technology to massively upgrade human intelligence is also a prerequisite. But that, on its own, may not be sufficient.

Compassion by those who possess overwhelming advantages over others is one of the special characteristics that Homo sapiens (along with a few other mammals) brings to this cold universe. It is what separates us from an asteroid or super-nova that doesn’t care whether it wipes us out.

Further, compassionate behavior is something most of us could agree on, and while it is often misinterpreted by some as a weakness, it is also what makes us human, and what most of us would want to contribute to future species.

If that is so, then let’s take the risk of being compassionate and put it into practice by launching overarching works that demonstrate the best of what we are.

For example, use molecular manufacturing and its predecessor nanotechnologies to eliminate the disease of aging, instead of treating the symptoms. That is what I personally have decided to focus on, but there are many other good examples out there, including synthesized meat that eliminates inhumane treatment of billions of animals, and cheap photovoltaic electricity that could slash our dependence on oil—and end wars over it.

Such works are not hard to identify. We just have to give them priority. Perhaps then we will seem less like our unwitting ancestors and more like enlightened guardians.

---

1. The Coming Technological Singularity: How to Survive in the Post-Human Era http://www-rohan.sdsu.edu/faculty/vinge/misc/singularity.html
2. http://www.wired.com/wired/archive/8.04/joy.html

© 2006 Douglas Mulhall. Reprinted with permission.

NASA's Dragonfly Project Demonstrates Robotic Satellite Assembly Critical to Future Space Infrastructure Development

Sept. 13, 2017

Original link:  https://www.nasa.gov/mission_pages/tdm/irma/nasas-dragonfly-project-demonstrates-robotic-satellite-assembly-critical-to-future-space.html

Shannon Ridinger
Marshall Space Flight Center, Huntsville, Ala.
256-544-0034
shannon.j.ridinger@nasa.gov

Last Updated: Sept. 13, 2017

Editor: Lee Mohon

This artist's rendering depicts Dragonfly assembling and

deploying large antenna reflectors on a satellite in Earth orbit.

Credits: NASA/SSL

A revolutionary NASA Technology Demonstration Mission project called Dragonfly, designed to enable robotic self-assembly of satellites in Earth orbit, has successfully completed its first major ground demonstration.

"Dragonfly has taken a crucial step toward rewriting the book on what we can do in Earth orbit -- and what that means for a robust space infrastructure," said John Lymer, chief architect of robotics and automation for Space Systems Loral (SSL) of Palo Alto, California, which leads the Dragonfly venture for NASA and conducted the testing at its Pasadena facility.

A lightweight robotic system with a dexterous 3.5-meter arm that's able to clamp down, carry items or operate controls -- from either end of the "limb" -- Dragonfly can install delicate satellite antenna, yet also assemble satellites too massive to be launched to space in their final flight-ready state.
These disassembled satellites may be stowed more efficiently or even launched in pieces via multiple flights, enabling mission planners to maximize cargo space and reduce mass. That shift would dramatically reduce launch costs and lead to less expensive, higher-performing satellites.

Dragonfly is one of three NASA tipping point projects seeking cutting-edge solutions under the umbrella of the TDM program's In-space Robotic Manufacturing and Assembly portfolio. These tipping point projects help the agency determine whether technologies have been sufficiently matured to pursue flight demonstrations or for infusion into future exploration missions.

"NASA relies on commercial innovation as exemplified by the Dragonfly team," said Trudy Kortes, TDM program executive at NASA Headquarters in Washington. "Transformative technologies such as these will, in time, lead to more affordable, safer human access to space and more efficient, longer-lasting satellites, probes and other space hardware. Today our future in space looks brighter and more robust than ever."

More about Dragonfly capabilities

During the August ground demonstration, Dragonfly's initial focus was the installation and reconfiguration of large antenna reflectors on a simulated geostationary satellite. The antennas are designed to focus the satellite signal to receivers on the ground. Additional demonstrations are planned through 2018 to further refine its processes and capabilities, including more fluid robotic arm movement and its ability to make even more precise reflector alignments.

Over time, the system will integrate 3-D printing technology enabling the automated manufacture of new antennae and even replacement reflectors as needed. Should a piece of hardware be damaged, or come to the end of its lifecycle, engineers could remotely remove and recycle the outdated component, replacing it with a new one.

"This is a major step forward in overhauling how we manage satellites in orbit," said Al Tadros, vice president for space infrastructure and civil space at SSL. He envisions a robust space infrastructure servicing industry, including commercial refueling platforms designed to service geostationary satellites and cargo missions to other planets.

Lymer concurred. "Being able to build what we need when we need it, change out hardware and components, even reconfigure old satellites for new tasks when they finish their initial missions, using affordable robotic technology," he said. "We're no longer at the mercy of launch costs and launch windows."

The Dragonfly project leverages a previous SSL study supported by Defense Advanced Research Projects Agency funding. Collaborating on the project are NASA's Langley Research Center in Hampton, Virginia; NASA's Ames Research Center in Moffett Field, California; Tethers Unlimited of Bothell, Washington; and SSL's affiliate MDA US Systems LLC of Pasadena, California. The project is sponsored by NASA's Space Technology Mission Directorate.

For more information about Dragonfly, visit:  http://www.sslmda.com/html/robotics_servicing.php

To learn more about NASA's Space Technology Mission Directorate, visit:  https://www.nasa.gov/spacetech

https://youtu.be/u_gYhG4L4QU

 

In this animation, the robotic Dragonfly system technology unpacks, assembles and deploys large antenna reflectors on a conventional satellite in geostationary Earth orbit. Dragonfly, a NASA Technology Demonstration Mission project pursuing advanced, in-space robotic manufacturing and assembly solutions, could reduce hardware and launch costs and in time lead to a robust space infrastructure and new solutions supporting exploration of our solar system.

Credits: Video courtesy of Space Systems Loral

 

Gene-centered view of evolution

From Wikipedia, the free encyclopedia

The gene-centered view of evolution, gene's eye view, gene selection theory, or selfish gene theory holds that adaptive evolution occurs through the differential survival of competing genes, increasing the allele frequency of those alleles whose phenotypic trait effects successfully promote their own propagation, with gene defined as "not just one single physical bit of DNA [but] all replicas of a particular bit of DNA distributed throughout the world". The proponents of this viewpoint argue that, since heritable information is passed from generation to generation almost exclusively by DNA, natural selection and evolution are best considered from the perspective of genes.

Proponents of the gene-centered viewpoint argue that it permits understanding of diverse phenomena such as altruism and intragenomic conflict that are otherwise difficult to explain.^[4][5]

The gene-centered view of evolution is a synthesis of the theory of evolution by natural selection, the particulate inheritance theory, and the non-transmission of acquired characters.^[6][7] It states that those alleles whose phenotypic effects successfully promote their own propagation will be favorably selected relative to their competitor alleles within the population. This process produces adaptations for the benefit of alleles that promote the reproductive success of the organism, or of other organisms containing the same allele (kin altruism and green-beard effects), or even its own propagation relative to the other genes within the same organism (intragenomic conflict).

The gene-centered perspective can be traced to the philosopher Henri Bergson who wrote, in his book, ‘Creative Evolution’ (1907)^[8]:

‘Life is like a current passing from germ to germ through the medium of a developed organism. It is as if the organism itself were only an excrescence, a bud caused to sprout by the former endeavouring to continue itself in a new germ’.

Overview

Ronald Fisher

 

John Maynard Smith

 

Richard Dawkins

The gene-centered view of evolution is a model for the evolution of social characteristics such as selfishness and altruism.

Acquired characteristics

The formulation of the central dogma of molecular biology was summarized by Maynard Smith:

If the central dogma is true, and if it is also true that nucleic acids are the only means whereby information is transmitted between generations, this has crucial implications for evolution. It would imply that all evolutionary novelty requires changes in nucleic acids, and that these changes – mutations – are essentially accidental and non-adaptive in nature. Changes elsewhere – in the egg cytoplasm, in materials transmitted through the placenta, in the mother's milk – might alter the development of the child, but, unless the changes were in nucleic acids, they would have no long-term evolutionary effects.

— Maynard Smith^[9]

The rejection of the inheritance of acquired characters, combined with Ronald Fisher the statistician, giving the subject a mathematical footing, and showing how Mendelian genetics was compatible with natural selection in his 1930 book The Genetical Theory of Natural Selection.^[10] J. B. S. Haldane, and Sewall Wright, paved the way to the formulation of the selfish-gene theory.^{[clarification needed]} For cases where environment can influence heredity, see epigenetics.^{[clarification needed]}

The gene as the unit of selection

The view of the gene as the unit of selection was developed mainly in the works of Richard Dawkins,^[11][12] W. D. Hamilton,^[13][14][15] Colin Pittendrigh^[16] and George C. Williams.^[17] It was mainly popularized and expanded by Dawkins in his book The Selfish Gene (1976).^[1]

According to Williams' 1966 book Adaptation and Natural Selection,

[t]he essence of the genetical theory of natural selection is a statistical bias in the relative rates of survival of alternatives (genes, individuals, etc.). The effectiveness of such bias in producing adaptation is contingent on the maintenance of certain quantitative relationships among the operative factors. One necessary condition is that the selected entity must have a high degree of permanence and a low rate of endogenous change, relative to the degree of bias (differences in selection coefficients).

— Williams,^[17] 1966, pp. 22–23

Williams argued that "[t]he natural selection of phenotypes cannot in itself produce cumulative change, because phenotypes are extremely temporary manifestations." Each phenotype is the unique product of the interaction between genome and environment. It does not matter how fit and fertile a phenotype is, it will eventually be destroyed and will never be duplicated.

Since 1954, it has been known that DNA is the main physical substrate to genetic information, and it is capable of high-fidelity replication through many generations. So, a particular gene coded in a nucleobase sequence of a lineage of replicated DNA molecules can have a high permanence and a low rate of endogenous change.^[18]

In normal sexual reproduction, an entire genome is the unique combination of father's and mother's chromosomes produced at the moment of fertilization. It is generally destroyed with its organism, because "meiosis and recombination destroy genotypes as surely as death."^[17] Only half of it is transmitted to each descendant due to independent segregation.

And the high prevalence of horizontal gene transfer in bacteria and archaea means that genomic combinations of these asexually reproducing groups are also transient in evolutionary time: "The traditional view, that prokaryotic evolution can be understood primarily in terms of clonal divergence and periodic selection, must be augmented to embrace gene exchange as a creative force."^[19][20]

The gene as an informational entity persists for an evolutionarily significant span of time through a lineage of many physical copies.^[2][21]

In his book River out of Eden, Dawkins coins the phrase God's utility function to explain his view on genes as units of selection. He uses this phrase as a synonym of the "meaning of life" or the "purpose of life". By rephrasing the word purpose in terms of what economists call a utility function, meaning "that which is maximized", Dawkins attempts to reverse-engineer the purpose in the mind of the Divine Engineer of Nature, or the utility function of god. Finally, Dawkins argues that it is a mistake to assume that an ecosystem or a species as a whole exists for a purpose. He writes that it is incorrect to suppose that individual organisms lead a meaningful life either; in nature, only genes have a utility function – to perpetuate their own existence with indifference to great sufferings inflicted upon the organisms they build, exploit and discard.

Organisms as vehicles

Genes are usually packed together inside a genome, which is itself contained inside an organism. Genes group together into genomes because "genetic replication makes use of energy and substrates that are supplied by the metabolic economy in much greater quantities than would be possible without a genetic division of labour."^[22] They build vehicles to promote their mutual interests of jumping into the next generation of vehicles. As Dawkins puts it, organisms are the "survival machines" of genes.^[1]

The phenotypic effect of a particular gene is contingent on its environment, including the fellow genes constituting with it the total genome. A gene never has a fixed effect, so how is it possible to speak of a gene for long legs? It is because of the phenotypic differences between alleles. One may say that one allele, all other things being equal or varying within certain limits, causes greater legs than its alternative. This difference enables the scrutiny of natural selection.

"A gene can have multiple phenotypic effects, each of which may be of positive, negative or neutral value. It is the net selective value of a gene's phenotypic effect that determines the fate of the gene."^[23] For instance, a gene can cause its bearer to have greater reproductive success at a young age, but also cause a greater likelihood of death at a later age. If the benefit outweighs the harm, averaged out over the individuals and environments in which the gene happens to occur, then phenotypes containing the gene will generally be positively selected and thus the abundance of that gene in the population will increase.

Even so, it becomes necessary to model the genes in combination with their vehicle as well as in combination with the vehicle's environment.

Selfish-gene theory

The selfish-gene theory of natural selection can be restated as follows:^[23]

Genes do not present themselves naked to the scrutiny of natural selection, instead they present their phenotypic effects. [...] Differences in genes give rise to differences in these phenotypic effects. Natural selection acts on the phenotypic differences and thereby on genes. Thus genes come to be represented in successive generations in proportion to the selective value of their phenotypic effects.

— Cronin, 1991, p. 60

The result is that "the prevalent genes in a sexual population must be those that, as a mean condition, through a large number of genotypes in a large number of situations, have had the most favourable phenotypic effects for their own replication."^[24] In other words, we expect selfish genes ("selfish" meaning that it promotes its own survival without necessarily promoting the survival of the organism, group or even species). This theory implies that adaptations are the phenotypic effects of genes to maximize their representation in future generations. An adaptation is maintained by selection if it promotes genetic survival directly, or else some subordinate goal that ultimately contributes to successful reproduction.

Individual altruism and genetic egoism

The gene is a unit of hereditary information that exists in many physical copies in the world, and which particular physical copy will be replicated and originate new copies does not matter from the gene's point of view.^[21] A selfish gene could be favored by selection by producing altruism among organisms containing it. The idea is summarized as follows:

If a gene copy confers a benefit B on another vehicle at cost C to its own vehicle, its costly action is strategically beneficial if pB > C, where p is the probability that a copy of the gene is present in the vehicle that benefits. Actions with substantial costs therefore require significant values of p. Two kinds of factors ensure high values of p: relatedness (kinship) and recognition (green beards).

— Haig,^[22] 1997, p. 288

A gene in a somatic cell of an individual may forego replication to promote the transmission of its copies in the germ line cells. It ensures the high value of p = 1 due to their constant contact and their common origin from the zygote.

The kin selection theory predicts that a gene may promote the recognition of kinship by historical continuity: a mammalian mother learns to identify her own offspring in the act of giving birth; a male preferentially directs resources to the offspring of mothers with whom he has copulated; the other chicks in a nest are siblings; and so on. The expected altruism between kin is calibrated by the value of p, also known as the coefficient of relatedness. For instance, an individual has a p = 1/2 in relation to his brother, and p = 1/8 to his cousin, so we would expect, ceteris paribus, greater altruism among brothers than among cousins. In this vein, geneticist J. B. S. Haldane famously joked, "Would I lay down my life to save my brother? No, but I would to save two brothers or eight cousins."^[25] However, examining the human propensity for altruism, kin selection theory seems incapable of explaining cross-familiar, cross-racial and even cross-species acts of kindness.

Green-beard effect

Green-beard effects gained their name from a thought-experiment of Richard Dawkins,^[1] who considered the possibility of a gene that caused its possessors to develop a green beard and to be nice to other green-bearded individuals. Since then, "green-beard effect" has come to refer to forms of genetic self-recognition in which a gene in one individual might direct benefits to other individuals that possess the gene. Such genes would be especially selfish, benefiting themselves regardless of the fates of their vehicles. After Dawkins predicted them, green-beard gene have been discovered in nature, such as Gp-9 in fire ants (Solenopsis invicta),^[26][27] csA in social amoeba (Dictyostelium discoideum),^[28] and FLO1 in budding yeast (Saccharomyces cerevisiae).^[29]

All kinds of altruism

Kindness

On the other hand, a single trait, group reciprocal kindness, is capable of explaining the vast majority of altruism that is generally accepted as "good" by modern societies. Imagine a green-bearding behavioral trait whose recognition does not depend on the recognition of some external feature such as beard color, but relies on recognition of the behavior itself. Imagine now that the behavior is altruistic. The success of such a trait in sufficiently intelligent and undeceived organisms is implicit. Moreover, the existence of such a trait predicts a tendency for kindness to unrelated organisms that are apparently kind, even if the organisms are of a completely different species. Moreover, the gene need not be exactly the same, so long as the effect is similar. Multiple versions of the gene—or even meme—would have virtually the same effect in a sort of symbiotic green-bearding cycle of altruism.

Deceit

Whenever recognition plays a role in evolution, so does deception. Just like the harmless lizard that has evolved a pattern that mimics its poisonous cousin and therefore tricks predators, the selfish creature may pretend to be kind by "growing a green beard" (whatever that green beard may be). Thus green-bearding and the selfish-gene theory also give rise to an explanation for the evolution of lies and deceit, characteristics that do not benefit the population as a whole.

Intragenomic conflict

As genes are capable of producing individual altruism, they are capable of producing conflict among genes inside the genome of one individual. This phenomenon is called intragenomic conflict and arises when one gene promotes its own replication in detriment to other genes in the genome. The classic example is segregation distorter genes that cheat during meiosis or gametogenesis and end up in more than half of the functional gametes. These genes persist even resulting in reduced fertility. Egbert Leigh compared the genome to "a parliament of genes: each acts in its own self-interest, but if its acts hurt the others, they will combine together to suppress it" to explain the relative low occurrence of intragenomic conflict.^[30]

Price equation

The Price equation is a covariance equation that is a mathematical description of evolution and natural selection. The Price equation was derived by George R. Price, working in London to rederive W. D. Hamilton's work on kin selection.

Advocates

Besides Richard Dawkins and George C. Williams, other biologists and philosophers have expanded and refined the selfish-gene theory, such as John Maynard Smith, George R. Price, Robert Trivers, David Haig, Helena Cronin, David Hull, Philip Kitcher, and Daniel C. Dennett.

Criticisms

The gene-centric view has been opposed by Ernst Mayr, Stephen Jay Gould, David Sloan Wilson, and philosopher Elliott Sober. An alternative, multilevel selection (MLS), has been advocated by E. O. Wilson, David Sloan Wilson, Sober, Richard E. Michod,^[31] and Samir Okasha.^[31]

Writing in the New York Review of Books, Gould has characterized the gene-centered perspective as confusing book-keeping with causality. Gould views selection as working on many levels, and has called attention to a hierarchical perspective of selection. Gould also called the claims of Selfish Gene "strict adaptationism", "ultra-Darwinism", and "Darwinian fundamentalism", describing them as excessively "reductionist". He saw the theory as leading to a simplistic "algorithmic" theory of evolution, or even to the re-introduction of a teleological principle.^[32] Mayr went so far as to say "Dawkins' basic theory of the gene being the object of evolution is totally non-Darwinian."^[33]

Gould also addressed the issue of selfish genes in his essay "Caring groups and selfish genes".^[34] Gould acknowledged that Dawkins was not imputing conscious action to genes, but simply using a shorthand metaphor commonly found in evolutionary writings. To Gould, the fatal flaw was that "no matter how much power Dawkins wishes to assign to genes, there is one thing that he cannot give them – direct visibility to natural selection."^[34] Rather, the unit of selection is the phenotype, not the genotype, because it is phenotypes that interact with the environment at the natural-selection interface. So, in Kim Sterelny's summation of Gould's view, "gene differences do not cause evolutionary changes in populations, they register those changes."^[35] Richard Dawkins replied to this criticism in a later book, The Extended Phenotype, that Gould confused particulate genetics with particulate embryology, stating that genes do "blend", as far as their effects on developing phenotypes are concerned, but that they do not blend as they replicate and recombine down the generations.^[12]

Since Gould's death in 2002, Niles Eldredge has continued with counter-arguments to gene-centered natural selection.^[36] Eldredge notes that in Dawkins' book A Devil's Chaplain, which was published just before Eldredge's book, "Richard Dawkins comments on what he sees as the main difference between his position and that of the late Stephen Jay Gould. He concludes that it is his own vision that genes play a causal role in evolution," while Gould (and Eldredge) "sees genes as passive recorders of what worked better than what".