Project
Association Theory
Initial Question
Why do organisms recognize kin?
When I entered graduate school in 2003, it was thought that kin recognition evolves in response to somatic parasites or "cheaters." The idea was that by restricting fusion or aggregation to kin, organisms would avoid being cheated by those that failed to act altruistically. In social amoebae, such cheaters would be those that avoided producing sterile stalk and only produced fertile spores. My goal was to isolate cheater mutants from a natural population and then show, in the laboratory, how they select for kin recognition. After searching for cheaters in a natural population of Dictyostelium discoideum, however, I found none! Yet, I found that D. discoideum, could recognize kin. By 2005, I started to wonder if there might be another explanation for kin recognition.
Approach
association Theory
Starting in late 2005, I began reading widely in hopes of developing a new model of kin recognition. I found that my best source of insight came from the detailed studies of botryllid ascidians. In mid 2008, I realized that costs of fusion could result from the mutual expression of selfish behaviors. By 2009, I had developed a new model of kin recognition. My model distinguished differential treatment (help/harm) and association preference (differentially entering the contexts for interactions). My model suggested that kin recognition could evolve for the purpose of association preference in response to differential treatment.
Predictions
Separate cues
Kin recognition involves a perception component and a cue component, and can evolve adaptively by modification of either component. A requirement for adaptive evolution of a trait is that it is genetically encoded. Most evidence suggests that genetic perception abilities arise by co-option of traits used for other purposes. Evidence suggests, however, that the cue component must evolve adaptively for the purpose of kin discrimination itself. They key feature that must be explained is the extreme genetic polymorphism of the cue, which ensures that allelic variants are unlikely to be shared with non-kin. Thus, sharing an allele means that individuals are likely related. In other words, the identity of the cue correlates with relatedness, and can be used as the basis for kin recognition.
My model showed that a requirement for the adaptive evolution of polymorphism is that separate cues are used for association preference as for differential treatment. My model showed that using different cues could be favored it cues used for differential treatment were not detectable at early stages of encounters. For example, a cue used for conflict might be detectable only at late stages of aggregation or fusion. Consequently, a behavior that used cues detectable earlier could be favored for avoiding conflict with non-kin. Given that separate cues are used, only a small amount of polymorphism of a cue used for differential treatment could exert a negative-frequency-dependent selective pressure for a cue used for association preference. I later found evidence that separate cues are used in bacteria, social amoebae, protochordates, cnidarians, fungi, plasmodial slime molds, arthropods, and vertebrates. If my inferences are correct, they are suggestive of the most successful genetic predictions in the history of biology because they apply across four levels of biological organization.
association Preference
In addition to explaining kin recognition, my model as yielded new predictions for the evolution of kin association preference. Previously, it was thought that organisms associate preferentially with kin in order to direct help to kin. No models showed how this is possible. Moreover, amphibian larvae and fish associate preferentially with kin but do not exhibit helping behavior. My model showed that such kin association preference could evolve in response to discriminatory harming behaviors, such as found in amphibians and fish. This also provided an explanation for association preference behaviors even in cooperative species, where harming behaviors are often prevalent.
fast-acting rejection
In a more recent article, I proposed that the evolution of fast-acting rejection responses in plasmodial slime molds and hydractiniid hydroids could have the same effect as switching cues. Under this hypothesis, multiple cues are used for discriminatory aggressive behaviors and some become fast-acting. The fast-acting aggressive behaviors then prevent fusion at early stages of encounters. Under this hypothesis, discriminatory aggressive behaviors morph into association preference behaviors that prevent fusion. Cues used for fast-acting "rejection" can then evolve polymorphism in response to those used for slow-acting aggressive behaviors.
Implications for social theory
Hamilton's rule
My model revealed that social theorists had long confused fundamental types of social behavior. For example, evolutionists had applied Hamilton's rule for altruism to association. This led to frequent violations of Hamilton's rule. In contrast, my theoretical framework, which I called "association theory," restricted Hamilton's rule to social actions, like altruism and selfishness, and did not apply it to association decisions. I showed how that this could prevent violations of Hamilton's rule. I also showed that it led to a reinterpretation of a textbook example of Hamilton's rule applied to turkeys.
Greenbeard effects
The basic idea of the "greenbeard effect" is that altruism can evolve among non-kin when a single gene codes for perception, a cue, and a discriminatory altruistic behavior. Social theorists took examples of aggregation and fusion among non-kin as evidence of the greenbeard effect. In reality, aggregation and fusion are examples of association behavior. Association differs from altruism because it merely provides the context for a social action. Association is also typically not cheatable in the same way as altruism. Social theorists therefore incorrectly inferred that greenbeard effects where they do not exist. Moreover, evolutionary models predict that greenbeards should be evolutionarily unstable and absent or rare in nature. My interpretation is consistent with this prediction.
Major transitions
Social theorists have split major transitions in biological individuality into three key steps: the origin, maintenance, and transformation of social groups. Typically, they apply Hamilton's rule to explain the origin of group living. They then typically downplay the importance of kin discrimination in major transitions. West et al. (2015) said that "within-group kin discrimination seems to have played a limited role in helping within species' major transitions." Incorporating the evolution of differential treatment and association preference into a theory of evolutionary transitions, however, suggest that they often involve the following steps: the the origin of association (group living), the evolution of differential treatment, the evolution of association preference, a switch in cues, and the evolution of kin recognition. This explains why in social insects, relatedness can drop in the absence of colonymate discrimination. It also explains why sedentary multicellular organisms that lack histocompatibility systems can fuse indiscriminately.
Inclusive fitness
My work on kin recognition suggested that there are three aspects of the inclusive fitness paradigm that were unfalsifiable. First, social theorists had long taken kin recognition as prima facie evidence of inclusive fitness. However, inclusive fitness theory had never yielded a falsifiable prediction for the evolution of kin recognition. It could therefore only confirm itself, explaining why it appeared so "successful."
Second, there was a large mathematical component of the inclusive fitness paradigm that sought to show that all complex design in nature was evidence of "inclusive fitness maximization." Although no models justified this paradigm, the continued search for a "Holy Grail" theorem kept the paradigm alive. In the end, however, nobody could agree what "inclusive fitness" is, much less derive a falsifiable empirical prediction from it.
Third, because inclusive fitness remained a murky concept, some authors generalized Hamilton’s rule to become what inclusive fitness was supposed to be: a general theorem of adaptation. This led to misapplications of Hamilton’s rule to phenomena that it does not explain. Selective reporting of apparent confirmations of Hamilton’s rule, in major journals and textbooks, gave the impression that Hamilton’s rule was rarely violated. Apparent violations were also dismissed with ad hoc arguments, while others were not reported. A consistent application of Hamilton’s rule, as determined by the inclusive-fitness paradigm, however, resulted in frequent violations. Because this was not emphasized due to selective reporting, again the paradigm could only confirm itself.
Returning social theory to Darwinian foundations
Organisms consist of thousands of genes, and of various genetic networks evolve in complex and idiosyncratic ways. To determine why a complex trait evolved, we must break it into simpler components and ask how the sub-component traits interact and became integrated into a complex whole. Some are resistant to such historical methods because they think that it means predictive power cannot be gained, in the same way it might come from a "fundamental theorem." It is thus important to keep in mind that fundamental theorems, like generalized versions of Hamilton's rule, have never yielded empirical predictions. In contrast, kin recognition has evolved many times independently, potentially through a common process. Predictive power can be gained by a stepwise historical model that yields predictions for diverse taxa.
Project
Natural Reward Theory
Initial Question
How might we extend Darwinism to explain macroevolution?
I had learned as an undergraduate that the key question for macroevolution was how factors like emergence and individuality define a level of selection. A major problem was whether species might reproduce in the same way as organisms. If so, natural selection might apply at the species level. In 2008, I realized that my model of kin recognition applied only to "organismal" levels. For example, it could help explain why unicellular, multinucleate, multicellular, and colonial organism remain genetically discrete. It would not explain why species remain genetically discrete. This suggested that there is a fundamental difference between organisms and species. This raised the question: how had Darwin justified this argument for within-species selection?
Approach
Reexamining core assumptions
I had learned from my textbooks that Darwin had defined the conditions for natural selection as (1) variation, (2) inheritance, and (3) differential reproductive success (or differential "fitness"). Thus, it seemed that natural selection might apply to any level that can reproduce ("has fitness"). Because species can seemingly reproduce with speciation and die with extinction, it seemed that natural selection might apply to the species level. Some argued that natural selection might also apply to clades, ecosystems, or entire universes!
If natural selection could so easily apply above the species level, why did Darwin not discuss this? Previously, I assumed that Darwin focused on within-species competition because this was relevant to the origin of species. But I know surmised that Darwin might have had a good reason to apply natural selection only within species!
The Struggle for existence
I thus reread Darwin with a questioning mind. I found that in addition to the three conditions mentioned above, Darwin had mentioned "the struggle for existence" as a requirement for natural selection. Contained in the "struggle for existence" was the idea that populations are limited by various "checks to increase." These could include those imposed by resource limitations, like food and water. However, they could also include limits on population growth imposed by predators and parasites. As a consequence of this assumption, only types within species would have the potential to competitively displace each other. Different species, relying on different resources and experiencing different "checks to increase" would often have the potential to coexist. Therefore, I realized that Darwin had used the "struggle for existence" to define the levels of selection.
Most evolutionists had dismissed Darwin's "struggle for existence" as an outdated slogan that he he used to get his theory across to a thick-headed audience. According to this view, Darwin's focus on death, competition, and "struggle" appealed to fashions of the day. It harkened Malthusian competition and Tennyson's (1850) "nature red in tooth and claw." Authors like Fisher, Dobzhansky, and Lewontin had derided Darwin's metaphor. It was thus largely eradicated from evolutionary discourse and textbooks, which is why nobody appreciated its logic.
But Darwin's argument made perfect sense. If two bacteria were growing in an abundance of nutrients, neither would completely displace the other until nutrients became limiting. In that case, if one started feeding on the metabolic waste product of the other, then the two strains would experience different checks to increase. Competitive displacement and natural selection would only occur within the strains. Likewise, two rainforest plants might look exactly the same, but experience different checks to increase imposed by predators owing to different chemical defenses. If one of these species had a slightly better metabolic ability it would not necessarily displace the other if predators checked it from increasing.
the Struggle for supremacy
By 2008, I was already thinking about the role of population growth in macroevolution. When I realized that Darwin had assumed that populations are checked from increasing, I realized that the key to extending Darwin's theory was to allow for transient population increases. I thus started developing a new theory based on "the struggle for supremacy," in which the first forms to escape checks to increase would expand in population, diversify, and be protected from invasion. In other words, the advantage of being first was in having a highly diversified and specialize suite of species, and the unit being rewarded would be the genetic system. Reading the literature, I found that the form of competition in macroevolution was not direct Darwinian competition as usually assumed, but instead usually required that an incumbent dwindle or go extinct before being replaced. This was consistent with the idea of an "incumbent advantage" and the notion of a "struggle for supremacy," which is also a term that Darwin once used in his unpublished manuscript Natural Selection.
natural reward
As I was developing this theory based on two different types of evolutionary struggle, I still thought that natural selection was the only non-random evolutionary force. I was increasingly having difficulty, however, keeping track of when natural selection was "selecting" alleles within species and when it was "rewarding" genetic systems. One day for convenience, I used "natural reward" to the effect of the incumbent advantage on favoring innovative forms of life. I soon realized that my study of kin recognition, and an increasing number of other studies, suggested that complex biological traits often originate for reasons that have nothing to do with the causes for success. This suggested that there was randomness at a level higher than mutation, in which the novel "inventions" produced by natural selection sometimes trigger biological expansions, thus becoming "innovations." I thus realize that "natural reward" might be a second non-random evolutionary force that acts upon the random variation of invention provided by natural selection (and other microevolutionary processes, like mutation and drift). I gradually came to believe, in contrast to the fabled Eureka! moment, that my "fudge" term of "natural reward" was a key to a new theory of macroevolution.
the success of the innovative
In 2019 and 2020, I outlined the theory of natural reward in two papers. My papers explained how that there had always been an ambiguity in Darwin's theory, about whether it explains relative or absolute progress. I argued that the ambiguity is in part what led to teleological theories, which applied natural selection to long-preserved units like species, clades, or immortal genes, or which argued that whole organisms are optimizing units. I then proposed the key components of the theory of natural reward and how it yields new insights on unsolved problems like the prevalence of sexual reproduction and the cause of punctuate equilibrium. I also argued that the theory helps define terms like progress and advancement, and that it predicts that life expands and becomes more innovative with time. I also reviewed studies of complex trait evolution that support my assumption of "randomness of invention." The theory of natural reward suggests that the broad-scale history of life is not "the survival of the fittest" but "the success of the innovative."
Practical implications
Darwin's theory of natural selection was deemed important because of its implications for everyday human life. It suggested that there is no progress in evolution, that man is just another animal, and that evolutionary theory gives no guidance for human ethics. It suggested that the history of life is summarized as "the survival of the fittest," where the "fittest" survives, reproduces, and eeks out its nearest competitors in the struggle for existence. Darwinism has sometimes been taken to justify "social Darwinism" and eugenics. Applied closely to scientific theories, it suggests that there is no scientific progress and that all truth is relative to dominant paradigms or historical power structures. Darwinism also meshes with a philosophy that suggests that human life is a zero-sum game, in which for one group to benefit, the other must be harmed. So its philosophy has been used to divide humanity into oppressor and oppressed categories.
The theory of natural that there is progress in evolution, that man is set apart his innovative capacity, and that evolutionary theory provides a guide for human ethics. It suggest that the history of life is summarized as “the success of the innovative,” and that nature rewards the most cooperative, creative, adaptable, and entrepreneurial forms of life. The theory of natural reward suggests that science progresses in explanatory power and ability to resolve anomalies. It suggests that the world is bountiful, and that innovation by one group of people can benefit others. It suggests that things are “good” that promote truth-seeking, cooperation, creativity, meritocracy, and prosperity, and “evil” those things that promote corruption, selfishness, conformity, nihilism, and poverty. The theory of natural reward also suggests that with innovation come creative destruction, such that a compassionate stance is to limit the negative effects of creative destruction by lifting up those who are marginalized and displaced. In that sense, the theory of natural reward provides a subtle approach to human ethics. It also has practical implications for making science more innovative (see below).
Project
Capitalistic science
Initial Question
Approach
what is wrong with science?
I went into graduate school with very ambitious goals. I wanted to use a study of kin recognition to gain insights on the "levels of selection" problem and macroevolution. To my surprise, I was led to the first fundamental advance in evolutionary theory since Darwin. I had succeeded beyond my wildest expectations, but this made it difficult to advance in my career. This was for five reasons. The first was that nearly all of the theorists in my field adopted the same paradigm, and were extremely conservative. A second was that the formed rings of mutual citation and funding, protecting the interests of their cronies during peer review. The third was that scientist within my field selectively reported confirmations of their unfalsifiable paradigm, thus exercising a level of corruption. The fourth was that my challenge to this paradigm caused them to censor my papers. A fifth was that scientists had become accustomed to evaluating others based on how expensive their research was, largely independently of the magnitude of discoveries. Science, I surmised, was overrun by the 5 C's: conservatism, cronyism, corruption, censorship, and costliness. This was also evidenced by numerous papers in the sociology of science.
Planned capitalism for science
Distilling the 5 C's down into a single issue, I realized that science was failing to innovate. This has since been born out by empirical studies (e.g., Park et al. 2023). My explanation for this, inspired by my macroevolutionary theory, is that science lacks rewards for innovation. Using a vast synthesis of economic literature, I have written a paper that proposes a way to complement the current funding structure of science with an auxiliary that is better geared for high-risk/high-reward research. The paper recognizes that the funding system of science is structured like a bureaucratic command economy and that an alternative must be structured more like free-market capitalism. Two key features that must be recovered are up-front funding for risky research and hindsight rewards for innovation. The problem, however is that many of the goods produced by science are not marketable and so the question is how to recover a capitalistic incentive structure. My solution is a prize based system based on crowdscoring, which encourages private investment in research and allows for a division of labor between researchers, entrepreneurs, and investors. I have argued that this system could be set up in a Web 3.0 ecosystem. I am currently revising a manuscript that proposes this funding system for science.
Many national command economies were also afflicted by the 5 C's, moreso than capitalistic economies. By creating an auxiliary capitalistic funding system for science, I believe this will also help eliminate the 5 C's. This is because if paradigms are more often challenged, then it less likely that false paradigms will persist for as long. There will then be less opportunity for the build up of questionable research practices that reinforce those paradigms. In my paper, I do not belabor these issues, because I do not want to distract from the greater message that we need an alternative system for funding risky research. This latter argument is not very controversial, and most scientists will be partial to it. Therefore, although I have communicated here some of the motivation for this line of research, via my own experiences, I am also cognizant of greater goals and the importance of being constructive.
I will finally add that I wanted to outline a capitalistic system for funding scientific research because I have surmised that what really defines capitalism is not a market per se, but the rewards for innovation and laws that promote a division of labor between inventors, entrepreneurs and investors that drives innovation. Showing how that a similar system could be created for scientific research, where the goods are non-appropriable, supports my argument. This will be included in my book on macroevolution.