Monday, October 3, 2011

Great evolution thinkers you should know about

If you follow a paleo diet, you follow a diet that aims to be consistent with evolution. This is a theory that has undergone major changes and additions since Alfred Russel Wallace and Charles Darwin proposed it in the 1800s. Wallace proposed it first, by the way, even though Darwin’s proposal was much more elaborate and supported by evidence. Darwin acknowledged Wallace's precedence, but received most of the credit for the theory anyway.

(Alfred Russel Wallace; source: Wikipedia)

What many people who describe themselves as paleo do not seem to know is how the theory found its footing. The original Wallace-Darwin theory (a.k.a. Darwin’s theory) had some major problems, notably the idea of blending inheritance (e.g., blue eye + brown eye = somewhere in between), which led it to be largely dismissed until the early 1900s. Ironically, it was the work of a Catholic priest that provided the foundation on which the theory of evolution would find its footing, and evolve into the grand theory that it is today. We are talking about Gregor Johann Mendel.

Much of the subsequent work that led to our current understanding of evolution sought to unify the theory of genetics, pioneered by Mendel, with the basic principles proposed as part of the Wallace-Darwin theory of evolution. That is where major progress was made. The evolution thinkers below are some of the major contributors to that progress.

Ronald A. Fisher. English statistician who proposed key elements of a genetic theory of natural selection in the 1910s, 1920s and 1930s. Fisher showed that the inheritance of discrete traits (e.g., flower color) described by Gregor Mendel has the same basis as the inheritance of continuous traits (e.g., human height) described by Francis Galton. He is credited, together with John B.S. Haldane and Sewall G. Wright, with setting the foundations for the development of the field of population genetics. In population genetics the concepts and principles of the theories of evolution (e.g., inheritance and natural selection of traits) and genetics (e.g., genes and alleles) have been integrated and mathematically formalized.

John B.S. Haldane. English geneticist who, together with Ronald A. Fisher and Sewall G. Wright, is credited with setting the foundations for the development of the field of population genetics. Much of his research was conducted in the 1920s and 1930s. Particularly noteworthy is the work by Haldane through which he mathematically modeled and explained the interactions between natural selection, mutation, and migration. He is also known for what is often referred to as Haldane’s principle, which explains the direction of the evolution of many species’ traits based on the body size of the organisms of the species. Haldane’s mathematical formulations also explained the rapid spread of traits observed in some actual populations of organisms, such as the increase in frequency of dark-colored moths from 2% to 95% in a little less than 50 years as a response to the spread of industrial soot in England in the late 1800s.

Sewall G. Wright. American geneticist and statistician who, together with Ronald A. Fisher and John B.S. Haldane, is credited with setting the foundations for the development of the field of population genetics. As with Fisher and Haldane, much of his original and most influential research was conducted in the 1920s and 1930s. He is believed to have discovered the inbreeding coefficient, related to the occurrence of identical genes in different individuals, and to have pioneered methods for the calculation of gene frequencies among populations of organisms. The development of the notion of genetic drift, where some of a population’s traits result from random genetic changes instead of selection, is often associated with him. Wright is also considered to be one of pioneers of the development of the statistical method known as path analysis.

Theodosius G. Dobzhansky. Ukrainian-American geneticist and evolutionary biologist who migrated to the United States in the late 1920s, and is believed to have been one of the main architects of the modern evolutionary synthesis. Much of his original research was conducted in the 1930s and 1940s. In the 1930s he published one of the pillars of the modern synthesis, a book titled Genetics and the Origin of Species. The modern evolutionary synthesis is closely linked with the emergence of the field of population genetics, and is associated with the integration of various ideas and predictions from the fields of evolution and genetics. In spite of Dobzhansky’s devotion to religious principles, he strongly defended Darwinian evolution against modern creationism. The title of a famous essay written by him is often cited in modern debates between evolutionists and creationists regarding the teaching of evolution in high schools: Nothing in Biology Makes Sense Except in the Light of Evolution.

Ernst W. Mayr. German taxonomist and ornithologist who spent most of his life in the United States, and is believed, like Theodosius G. Dobzhansky, to have been one of the main architects of the modern evolutionary synthesis. Mayr is credited with the development in the 1940s of the most widely accepted definition of species today, that of a group of organisms that are capable of interbreeding and producing fertile offspring. At that time organisms that looked alike were generally categorized as being part of the same species. Mayr served as a faculty member at Harvard University for many years, where he also served as the director of the Museum of Comparative Zoology. He lived to the age of 100 years, and was one of the most prolific scholars ever in the field of evolutionary biology. Unlike many evolution theorists, he was very critical of the use of mathematical approaches to the understanding of evolutionary phenomena.

William D. Hamilton. English evolutionary biologist (born in Egypt) widely considered one of the greatest evolution theorists of the 20th Century. Hamilton conducted pioneering research based on the gene-centric view of evolution, also know as the “selfish gene” perspective, which is based on the notion that the unit of natural selection is the gene and not the organism that carries the gene. His research conducted in the 1960s set the foundations for using this notion to understand social behavior among animals. The notion that the unit of natural selection is the gene forms the basis of the theory of kin selection, which explains why organisms often will instinctively behave in ways that will maximize the reproductive success of relatives, sometimes to the detriment of their own reproductive success (e.g., worker ants in an ant colony).

George C. Williams. American evolutionary biologist believed to have been a co-developer in the 1960s, together with William D. Hamilton, of the gene-centric view of evolution. This view is based on the notion that the unit of natural selection is the gene, and not the organism that carries the gene or a group of organisms that happens to share the gene. Williams is also known for his pioneering work on the evolution of sex as a driver of genetic variation, without which a species would adapt more slowly in response to environmental pressures, in many cases becoming extinct. He is also known for suggesting possible uses of human evolution knowledge in the field of medicine.

Motoo Kimura. Japanese evolutionary biologist known for proposing the neutral theory of molecular evolution in the 1960s. In this theory Kimura argued that one of the main forces in evolution is genetic drift, a stochastic process that alters the frequency of genotypes in a population in a non-deterministic way. Kimura is widely known for his innovative use of a class of partial differential equations, namely diffusion equations, to calculate the effect of natural selection and genetic drift on the fixation of genotypes. He has developed widely used equations to calculate the probability of fixation of genotypes that code for certain phenotypic traits due to genetic drift and natural selection.

George R. Price. American geneticist known for refining in the 1970s the mathematical formalizations developed by Ronald A. Fisher and William D. Hamilton, and thus making significant contributions to the development of the field of population genetics. He developed the famous Price Equation, which has found widespread use in evolutionary theorizing. Price is also known for introducing, together with John Maynard Smith, the concept of evolutionary stable strategy (ESS). The EES notion itself builds on the Nash Equilibrium, named after its developer John Forbes Nash (portrayed in the popular Hollywood film A Beautiful Mind). The concept of EES explains why certain evolved traits spread and become fixed in a population.

John Maynard Smith. English evolutionary biologist and geneticist credited with several innovative applications of game theory (which is not actually a theory, but an applied branch of mathematics) in the 1970s to the understanding of biological evolution. Maynard Smith is also known for introducing, together with George R. Price, the concept of evolutionary stable strategy (EES). As noted above, the EES notion builds on the Nash Equilibrium, and explains why certain evolved traits spread and become fixed in a population. The pioneering work by John Maynard Smith has led to the emergence of a new field of research within evolutionary biology known as evolutionary game theory.

Edward O. Wilson. American evolutionary biologist and naturalist who coined the term “sociobiology” in the 1970s to refer to the systematic study of the biological foundations of social behavior of animals, including humans. Wilson was one of the first evolutionary biologists to convincingly argue that human mental mechanisms are shaped as much by our genes as they are by the environment that surrounds us, setting the stage for the emergence of the field of evolutionary psychology. Many of Wilson’s theoretical contributions in the area of sociobiology are very general, and apply not only to humans but also to other species. Wilson has been acknowledged as one of the foremost experts in the study of ants’ and other insects’ social organizations. He is also known for his efforts to preserve earth’s environment.

Amotz Zahavi. Israeli evolutionary biologist best known for his widely cited handicap principle, proposed in the 1970s, which explains the evolution of fitness signaling traits that appear to be detrimental to the reproductive fitness of an organism. Zahavi argued that traits evolved to signal the fitness status of an organism must be costly in order to the reliable. An example is the large and brightly colored trains evolved by the males of the peacock species, which signal good health to the females of the species. The male peacock’s train makes it more vulnerable to predators, and as such is a costly indicator of survival success. Traits used for this type of signaling are often referred to as Zahavian traits.

Robert L. Trivers. American evolutionary biologist and anthropologist who proposed several influential theories in the 1970s, including the theories of reciprocal altruism, parental investment, and parent-offspring conflict. Trivers is considered to be one of the most influential living evolutionary theorists, and is a very active researcher and speaker. His most recent focus is on the study of body symmetry and its relationship with various traits that are hypothesized to have been evolved in our ancestral past. Trivers’s theories often explain phenomena that are observed in nature but are not easily understood based on traditional evolutionary thinking, and in some cases appear contradictory with that thinking. Reciprocal altruism, for example, is a phenomenon that is widely observed in nature and involves one organism benefiting another not genetically related organism, without any immediate gain to the organism (e.g., vampire bats regurgitating blood to feed non-kin).

There are many other more recent contributors that could arguably be included in the list above. Much recent progress has been made in interdisciplinary fields that could be seen as new fields of research inspired in evolutionary ideas. One such field is that of evolutionary psychology, which has emerged in the 1980s. New theoretical contributions tend to take some time to be recognized though, as will be the case with ideas coming off these new fields, because new theoretical contributions are invariably somewhat flawed and/or incomplete when they are originally proposed.


Mike said...

No doubt all great thinkers in some regards. Way too many of them were into eugenics. That's why you always have to pay attention to what a person is saying and not who. Nevertheless, these men all made some great contributions.

Ned Kock said...

Hi Mike. While I think these thinkers should be recognized for their great ideas, we should always be careful about the “argumentum ad verecundiam”, better known as the “argument from authority” fallacy.

Many of these great thinkers had ideas outside their sphere of competence that were at best wrong and at worst repulsive. Still some of those ideas were supported based on the “argument from authority” fallacy. Eugenics, as you said, is a common theme in attempts by these evolutionary biologists to contribute to the area of public policy.

Darwin referred to hunter-gatherers generally as “savages”, and considered them to be inferior races in evolutionary terms. One of Fisher’s main motivations for doing evolutionary biology research was admittedly the “disappearance of the higher classes in Britain” (peerages, which he saw as superior). Hamilton reportedly promoted infanticide à la the Spartans … and the list goes on.

Ned Kock said...

Btw, propensity toward odd ideas in general is also a theme among the great thinkers listed here. This could be seen as a “curse” of evolutionary thinking, but I think it is due to another problem. The problem could also be seen as a “curse”, associated with high levels of creative intelligence, and also with scoring very high in one of the big five personality traits – openness.

In short, great thinkers seem to be disproportionately more likely to suffer from mental disorders, which tend to distort their thinking. And this is in part due to evolution itself! I promise, one of my next posts will be about this, and it is related to the notion of costly traits evolved through sexual selection.

One of the most compelling and tragic examples of this “curse” is the case of George Price, perhaps the most creative of all the thinkers listed on this post. See this book:

Mike said...

Hi Ned,

I've put that book into my reading list, thank you.

I don't really have much to add because I completely agree with everything you just said.

David Isaak said...

I'm not sure it's quite correct to to say that Wallace had precedence over Darwin or that Darwin acknowledged it. Darwin had been working on the problem, and had written extensively about it (without publication) for two decades befoe he received Wallace's letter..and it was a letter, not a publication. So at the time they first made contact, both of them had been working on the idea and neither had published. Darwin always made sure that Wallace was given credit, but as far as I know he never claimed Wallace had precedence.

My favorite of the bunch on your list is the iconic JBS Haldane.

Haldane was once interviewed on the radio by a clergyman who asked what a lifetime of study of biology had taught Haldane about the nature of The Creator.

Haldane replied, "He has an inordinate fondness for beetles."

(Coleoptera contains more species than any other order of insects, and a large fraction of all the known species of animate life forms.)

Ned Kock said...

Haldane was probably also the most colorful, and certainly not one to resort to BS, in spite of his name ;-)

Ned Kock said...
This comment has been removed by the author.
Ned Kock said...

The Wallace-Darwin episode is debatable, because what “publication” meant at the time was very different from what it means today.

One thing is for sure though, animal breeders knew about the basic principles of evolution, well before the theory of evolution was formally proposed.

This includes George Washington:

Ned Kock said...

The first publication by Wallace related to evolution was, I believe, titled “On the Law which has Regulated the Introduction of New Species”. It was first published in the “Annals and Magazine of Natural History”, in September 1855.

It is available from this link:

Ned Kock said...

From a note at the end of the paper linked above:

“This paper was written three years before Wallace sent that fateful communication (S43) to Darwin in which he proposed that selective forces acting on heritable variation was a sufficient explanation for evolutionary change. Reading the present paper in the year 2000 (and with the benefit of hindsight, of course) it is difficult to believe that Wallace had not worked it all out at least three years earlier than generally admitted.”

David Isaak said...

The Darwin-Wallace precedence thing has been through a lot of scrutiny. I agree 'publication' didn't mean much at the time. The big competition of the day wasn't over theory in any case, but rather species discovery and who got to bestow species names. Some of the species hunters were positively nutty on the issue of priority*. (There's a fun book called The Species Seekers on the topic.)

*[Including the paleontologists; there were furious rivalries over fossil finds.]

There was a back-and-forth on the Darwin-Wallace issue in The Guardian a few years ago. This brief letter I think outlines the history fairly well, and also has links in the sidebars to other bits of the exchange

In any case, neither Darwin nor Wallace were attempting to discredit the other. Unlike, say, Leibniz and Newton, Darwin and Wallace weren't rivals, and both were careful to give credit to the other.

Certainly Wallace never expressed any complaints about the fact they were presented as co-discoverers.

In any case, as you mention, humans understood breeding and crossing already, and evolution was already a widespread idea. But it was a pretty flaky, fuzzy, semi-mystical idea until Wallace and Darwin.

(Actually, Wallace, like Newton, had a mystical streak, too.)

gwarm said...

Take a look at this Fibonacci sequence explained through a beautiful video
'Applications include computer algorithms such as the Fibonacci search technique and the Fibonacci heap data structure, and graphs called Fibonacci cubes used for interconnecting parallel and distributed systems. They also appear in biological settings,[6] such as branching in trees, arrangement of leaves on a stem, the fruit spouts of a pineapple,[7] the flowering of artichoke, an uncurling fern and the arrangement of a pine cone.[8]' wiki

Saw this video linked here

Ned Kock said...

Beautiful clip gwarm, thanks.

xl pharmacy said...

It is impressive the amount of diets that it is available out side but it is good because one can work with you and other don't

Unknown said...

Hi Ned. I saw the following and wondered if you might like to see them. Perhaps you could comment in a future blog post.