Friday, April 16, 2010

More on Evolution and Human Free Will


Every summer I teach a seminar course at Cornell in which we examine the historical, philosophical, religious, and scientific implications of evolutionary theory. This summer our seminar course will once again consider the question: Is free will an illusion?

On the 15th of July, 1838, Charles Darwin began a notebook which he labeled as “M”, in which he intended to write down his correspondence, discoveries, musings, and speculations on “Metaphysics on Morals and Speculations on Expression”. On page 27 of that notebook, he wrote
“…one doubts existence of free will every action determined by hereditary constitution, example of others or teaching of others. (…man…probably the only [animal] affected by various knowledge which is not heredetary & instinctive) & the others are learnt, what they teach by the same means & therefore properly no free will. [Emphasis added]

In his private musing on the question of free will, Darwin came to the conclusion that human free will is an illusion, and that all of our actions (and, by extension, our thoughts and intentions) are the result of our “hereditary constitution” and “the example…or teaching of others.”

Some evolutionary biologists, notably William Provine of Cornell University, have followed Darwin’s lead and asserted that human free will is an illusion. Most philosophers disagree, asserting that free will is the principle difference between humans and non-human animals. Many Christian theologians go further, asserting that free will is the foundation of all human action, without which no rational ethics or theology is possible.

In our seminar course this summer we will take up this debate by considering two alternative hypotheses: (1) that human free will is real and can provide a basis for our morals and ethics, or (2) that human free will is an illusion, the capacity for which is a product of the same evolutionary processes that have shaped our anatomical and behavioral adaptations. Included in this debate will be an extended consideration of the hypothesis that the capacity for ethical decision making is an evolutionary adaptation that has evolved by natural selection. We will read from some of the leading authors on both sides of the subject, including George Ainslie, Daniel Dennett, Robert Kane, William Provine, Daniel Wegner, and Edward O. Wilson. Our intent will be to sort out the various issues at play, and to come to clarity on how those issues can be integrated into a perspective of the interplay between philosophy and the natural sciences.

Here are some particulars for the course:

INTENDED AUDIENCE: This course is intended primarily for students in biology, history, philosophy, religious studies, and science & technology studies. The approach will be interdisciplinary, and the format will consist of in-depth readings across the disciplines and discussion of the issues raised by such readings.

PREREQUISITES: None, although a knowledge of general evolutionary theory, evolutionary psychology, sociobiology, and the philosophy of human free will would be useful.

DAYS, TIMES, & PLACES: The course will meet on Tuesday and Thursday evenings from 6:00 to 9:00 PM in Mudd Hall, Room 409 (The Whittaker Seminar Room), beginning on Tuesday 29 June 2010 and ending on Thursday 5 August 2010.

CREDIT & GRADES: The course will be offered for 4 hours of credit, regardless of which course listing students choose to register for. Unless otherwise noted, course credit in BIOEE 4670 / BSOC 4471 can be used to fulfill biology/science distribution requirements and HIST 4150 / STS 4471 can be used to fulfill humanities distribution requirements (check with your college registrar's office for more information). Letter grades for this course will be based on the quality of written work on original research papers written by students, plus participation in class discussion. All participants must be registered in the Cornell Six-Week Summer Session to attend class meetings and receive credit for the course (click here for for more information and to enroll for this course). Registration will be limited to the first 18 students who enroll for credit.

REQUIRED TEXTS:

Ainslie, G. (2008) Breakdown of Will, Cambridge University Press, ISBN: 0521596947 (paperback: $34.99), 272 pages.

Dennett, D. (2004) Freedom Evolves, Penguin Books, ISBN: 0142003840 (paperback: $17.00), 368 pages.

Kane, R. (2005) A Contemporary Introduction to Free Will, Oxford University Press (USA), ISBN: 019514970X (paperback: $19.95), 208 pages.

Wegner, D. (2003) The Illusion of Conscious Will, MIT Press, ISBN-10: 0262731622 (paperback: $21.95), 419 pages.

Wilson, E. O. (2004) On Human Nature (Revised Edition), Harvard University Press, ISBN: 0674016386 (paperback: $22.00), 284 pages.

OPTIONAL TEXTS:

Darwin, Charles (E. O. Wilson, ed.) (2006) From So Simple a Beginning: Darwin's Four Great Books. W. W. Norton, ISBN-10: 0393061345 (hardcover, $39.95), 1,706 pages. Available online here.

Fisher, J., Kane, R., Pereboom, D., & Vargas, M. (2007) Four Views on Free Will, Wiley-Blackwell, ISBN: 1405134860 (paperback: $33.95), 240 pages.

Kane, R. (2001) Free Will (Blackwell Readings in Philosophy), Wiley-Blackwell, ISBN: 0631221026 (paperback: $33.95), 328 pages.

Wilson, E. O. (2000) Sociobiology: The New Synthesis (25th Anniversary Edition), Belknap Press, ISBN: 0674002350 (paperback: $44.00), 720 pages

Our summer seminar course is always fascinating, and often quite controversial (see this and this). Over the years we have explored many of the implications of Darwin's theory, and the participants have always found our discussions (perhaps they should be called "debates") enlightening. As always, the intent is not necessarily to reach unanimity, but rather for each participant to come to clarity on where they stand on the issues and to be able to defend that stance using evidence and rational argument.

So, please consider taking our seminar on free will this summer - the choice is yours!

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As always, comments, criticisms, and suggestions are warmly welcomed!

--Allen

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Thursday, November 19, 2009

The Modern Synthesis is Dead - Long Live the Evolving Synthesis!


It has been almost exactly a century and a half since Darwin's Origin of Species was first published, and half a century since the conference at the University of Chicago where the "triumph" of the "modern evolutionary synthesis" was celebrated. So, isn't it a little odd that some well-respected scientists and historians of science are proclaiming in this celebratory year that the modern evolutionary synthesis is dead?

For example, Eugene Koonin, senior investigator at the National Center for Biotechnology Information, National Library of Medicine, and National Institutes of Health in Bethesda, Maryland, has published two essays on the current status of the "modern evolutionary synthesis":
The Origin at 150: Is a new evolutionary synthesis in sight?
Trends in Genetics, 25(11), November 2009, pp. 473-475.

Abstract: The 200th anniversary of Charles Darwin and the 150th jubilee of the On the Origin of Species could prompt a new look at evolutionary biology. The 1959 Origin centennial was marked by the consolidation of the modern synthesis. The edifice of the modern synthesis has crumbled, apparently, beyond repair. The hallmark of the Darwinian discourse of 2009 is the plurality of evolutionary processes and patterns. Nevertheless, glimpses of a new synthesis might be discernible in emerging universals of evolution.

and
Darwinian evolution in the light of genomics.
Nucleic Acids Research, 37(4), 2009, pp. 1011-1034.

ABSTRACT: Comparative genomics and systems biology offer unprecedented opportunities for testing central tenets of evolutionary biology formulated by Darwin in the Origin of Species in 1859 and expanded in the Modern Synthesis 100 years later. Evolutionary-genomic studies show that natural selection is only one of the forces that shape genome evolution and is not quantitatively dominant, whereas non-adaptive processes are much more prominent than previously suspected. Major contributions of horizontal gene transfer and diverse selfish genetic elements to genome evolution undermine the Tree of Life concept. An adequate depiction of evolution requires the more complex concept of a network or 'forest' of life. There is no consistent tendency of evolution towards increased genomic complexity, and when complexity increases, this appears to be a nonadaptive consequence of evolution under weak purifying selection rather than an adaptation. Several universals of genome evolution were discovered including the invariant distributions of evolutionary rates among orthologous genes from diverse genomes and of paralogous gene family sizes, and the negative correlation between gene expression level and sequence evolution rate. Simple, non-adaptive models of evolution explain some of these universals, suggesting that a new synthesis of evolutionary biology might become feasible in a not so remote future.

A big deal, right? Well, not really. Will Provine and I have been saying that “the modern evolutionary synthesis is dead” for years. Indeed, Will Provine coined the phrase “the hardening of the synthesis” to describe the narrowing of focus in evolutionary theory during the first half of the 20th century to concepts entirely reducible to mathematical models, especially theoretical population genetics.

Ironically, Dr. John Sanford and Dr. William Dembski (among others in the ID camp) have not moved beyond this narrow focus on theoretical population genetics, and so have apparently missed the fact that evolutionary biology has evolved far beyond the narrow theoretical focus of the mid-20th century. Some ID supporters have also suggested that Dr. Koonin might be taking a “big career risk” in stating the obvious. I don't think so. On the contrary, what Dr. Koonin has pointed out is that evolutionary biology today is broader, more generally applicable, and less narrowly focused than at any time since the publication of the Origin of Species 150 years ago. Being an evolutionary biologist today is like being a physicist in 1905 — a whole new world of theoretical and practical empirical research is opening up, with new discoveries being made every day.

As just one example, Kyoto-prize-winning evolutionary biologists Peter and Rosemary Grant have reported on something that Darwin could only speculate about: the systematic empirical documentation of the “origin” of a new species (reported here yesterday). Creationists have of course moved the goalposts, arguing that they accepted all along that new species could arise from existing ones, it’s just microevolution, which of course everyone accepts. This, despite the fact that speciation has always been considered to be the first (and perhaps most important) stage in macroevolution, and that less than two decades ago creationists were confidently stating that “true” speciation had not only never been observed, it couldn’t ever be observed because it can’t happen.

Now the leaders of the ID movement — people like Dr. Michael Behe and Dr. William Dembski — publicly state that they fully accept that descent with modification from common ancestors (i.e. evolution) has happened, that microevolution (i.e. natural selection, sexual selection, and genetic drift) are also fully supported by the evidence, and that the “real” focus of disagreement is over the “engines of variation” that produce the raw material upon which the “engines of evolution” operate. They’ve come a long way, but they’ve missed the parade by a couple of decades. So it goes…

I would say that Dr. Koonin's essays on where evolutionary biology is today are quite close to the the mark. The concept of natural selection as the foundation of evolutionary change has been largely superseded, mostly through the work of Motoo Kimura, Tomoko Ohta, and others, who have shown both theoretically and empirically that natural selection has little or no effect on the vast majority of the genomes of most living organisms.

However, ID supporters should find this sea change in evolutionary biology to be cold comfort. The overall effect of the advances in our understanding of how genomes and phenotypes change over time has had the same effect on evolutionary theory that the rise of quantum mechanics had on classical physics. Einstein famously asserted that “God does not play dice”, but a century of physics research has shown him to be more wrong about how the universe works at the quantum level than ever.

The same is true for the “evolving synthesis”. Rather than revert to a neo-Paleyan paradigm (as proposed by Behe, Dembski, and their supporters), evolutionary biology has gone in the opposite direction, the same direction that quantum mechanics has taken. According to the “modern synthesis” of the last century, the genome was “homeostatic”, “organized”, and “regulated” primarily by natural selection. Sure there were purely random processes also going on (such as genetic drift), but most evolutionary change was both adaptive and coherent over time.

Here's what Dr. Koonin writes (see above):
"There is no consistent tendency of evolution towards increased genomic complexity, and when complexity increases, this appears to be a nonadaptive consequence of evolution under weak purifying selection rather than an adaptation."

Kimura, Ohta, Jukes, and Crow dropped a monkey wrench into the "engine" at the heart of the modern synthesis — natural selection — and then Gould and Lewontin finished the job with their famous paper on “the spandrels of San Marco and the Panglossian paradigm”. The rise of evo-devo over the past two decades has laid the groundwork for a completely new and empirically testable theory of macroevolution, a theory that is currently facilitating exponential progress in our understanding of how major evolutionary transitions happen. And iconoclasts like Lynn Margulis, Eva Jablonka, Marian Lamb, Mary Jane West-Eberhard, and David Sloan Wilson are rapidly overturning our understanding of how evolutionary change happens at all levels, and how it is inherited.

So, as I have said many times before, when ID supporters set their sights on “neo-Darwinism” as a target for criticism, they set their sights on a model that has been all but abandoned. The carnival has moved on and ID supporters are fighting battles that evolutionary biologists left behind a half century and more ago.

And so, on this 150th anniversary of the publication of Darwin's Origin of Species, evolutionary biologists can raise a frosty glass and say


The modern synthesis is dead — long live the evolving synthesis!


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As always, comments, criticisms, and suggestions are warmly welcomed!

--Allen

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Sunday, November 08, 2009

The Darwinian Revolutions Video Series


The Darwinian Revolutions

An online video lecture series
in honor of the 150th anniversary
of the original publication of
Charles Darwin's Origin of Species


Produced by:
The Cybertower at Cornell University

Written, directed and narrated by:
Allen MacNeill, Senior Lecturer
The Biology Learning Skills Center
Ecology & Evolutionary Biology

Videography by:
Dina Banning

Sound Engineering by:
Colbert McClennan

Technical Direction by:
Becky Lane

Videotaped at:
The Museum of the Earth
The Paleontological Research Institution
Ithaca, New York

Voiceover Narration Recorded at:
Fall Creek Studios
1201 North Tioga Street
Ithaca, New York

Images Obtained at:
WikiMedia Commons
Stebbins/Simpson/Dobzhanky photo credit: Martin Tracey

Galapagos Video Credit:
Prof. William Provine

It's finally done! After more than a year of meetings, writing, image acquisition, videotaping, sound recording, editing, revising, captioning, and (most of all) thinking, our video series on the Darwinian revolutions is now online!

This series of six online videos is a brief introduction to Darwin's theory of evolution by natural selection and its implications. Here is a brief synopsis of the six episodes (click on each episode title to go to the linked video):


Episode One: Darwinian Revolutions
We begin with an overview of the series, which has been released to coincide with the 150th anniversary of the publication of Darwin's Origin of Species. Darwin's theory of evolution by natural selection revolutionized both the biological sciences and our understanding of ourselves and the world around us. In this episode we learn that Darwin's theory has itself evolved in the 150 years since it was published. We also learn that Darwin actually presented two theories:
• a theory of descent with modification from common ancestors, and
• the theory of natural selection, Darwin's mechanism for evolution.


Episode Two: Evolutionary Ancestors
Beginning with an overview of Darwin's predecessors, we learn how the idea of evolution by natural means alone goes back more than two thousand years, to ancient Greece and Rome. Democritus of Abdera first proposed the "ground rules" for naturalist evolution, which were later extended by the Roman poet and philosopher, Lucretius. However, these early naturalistic theories were eclipsed for almost two millennia by the ideas of Plato and Aristotle.


Episode Three: Lamarck's Theory
In the 19th century, Jean Baptiste Lamarck set the stage for Darwin's monumental achievement with his Philosophie Zoologique (published in 1809), which advanced a theory of evolution by means of the inheritance of acquired characteristics. Lamarck's theory was the first theory of evolution to include a testable mechanism for evolutionary change — the inheritance of acquired characteristics — and provides a useful comparison with Darwin's theory.


Episode Four: One Long Argument
Darwin, whose academic training at Cambridge University was in Anglican theology, became an acclaimed naturalist and science writer following the five-year voyage of HMS Beagle. Using the notes and specimens that he had collected during the voyage, Darwin spent twenty years refining his theory, first published in 1859, of evolution by natural selection.


Episode Five: Mendel and the Eclipse of Darwin
Darwin's theory of descent with modification was accepted by most scientists worldwide within ten years of its publication in 1859. However, his theory of natural selection was widely criticized, and by the turn of the 20th century was widely considered to be dead. However, the work of Gregor Mendel, who discovered the foundations of what we now call genetics, provided a mechanism by which Darwin's theory could be revived and expanded.


Episode Six: The Evolving Synthesis
In the final segment of this series, we visit the The Museum of the Earth in Ithaca, New York, whose director, Dr. Warren Allman, discusses the importance of such museums to the science of evolutionary biology. We also hear from Cornell professor William Provine, who discusses Darwin's work and its importance to the history and philosophy of biology. He tells us how Darwin's original theory of natural selection was integrated into the sciences of population genetics, ecology, physiology, paleontology, embryology, and botany, to produce a "modern synthesis" of evolutionary theory. Prof. Provine also tells us how the "modern synthesis" has continued to evolve, and that today is the most exciting time yet in the history of Darwin's scientific revolution.

This has been an exciting year: the 200th anniversary of the publication of Lamarck's Philosophy Zoologique, the 200th anniversary of the birth of Charles Darwin, and the 150th anniversary of the publication of Darwin's Origin of Species. There have been many events marking these anniversaries, and there will be many more. As Will Provine says, the theory of evolution is more dynamic, more exciting, more widely accepted, and more widely applied than at any time in the past century and a half. With the accelerating pace of discoveries in evolutionary biology and their applications in biology, medicine, psychology, economics, and even literature and art, the 21st century shows all indications of being what the founders of the "modern synthesis" called it back in 1959: the "century of Darwin" and his theory of evolution by natural selection.

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As always, comments, criticisms, and suggestions are warmly welcomed!

--Allen

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Tuesday, March 24, 2009

Intelligent Design is Boring


At a thread at another website discussing the idea that ID is "boring", an ID supporter wrote this:

"[ID] is boring to Will [Provine] I suspect (and to others for the same reason) because they rule out the possibility of an intelligent designer."

Actually, knowing Will Provine pretty well and hearing him say that ID is "boring" on several occasions, I can confidently state that the reason he finds it "boring" is that whenever something interesting in biology is discovered and somebody asks "Why is that thing the way it is?" Will hears most ID supporters answer "Goddidit". His opinion of ID is that it's a science-stopper because rather than suggesting new and interesting ways of trying to figure out how something came to be the way it is, he thinks that IDers simply throw up their hands and say "It's too complicated, so God / the Intelligent Designer must have done it".

Personally, I don't find ID boring for quite the same reason, as I don't always see ID supporters resorting to the "Goddidit" pseudoexplanation. No, the reason I tend to find most ID boring is it's relentlessly negative. That is, people like Michael Behe and William Dembski observe something marvelously complicated and say "That's Irreducibly Complex!" or "That's Complex Specified Information, so it couldn't have evolved via naturalistic means"...and then they leave it at that. No alternative means of creating the marvelously complicated thing is proposed (unless you credit Behe's "puff of smoke" pseudoargument).

Furthermore, I generally don't see ID supporters doing any original empirical research. In particular, I don't see any of them going out into the field (my favorite place to discover things) or into the lab and "getting down and dirty" with some biological phenomenon that they find absolutely fascinating.


My friend, Harry Greene (the world's authority on rattlesnakes) is my idea of a real scientist. He absolutely loves snakes, talks about them at the drop of a hat, has spent his entire professional life studying them in the field and in the lab, and has revolutionized our understanding of the ecology, ethology, and evolutionary biology of reptiles. To me, he's the epitome of an evolutionary biologist, because he has what we call "a feel for the organism" which goes far beyond simply studying it as an experimental subject.


And my friend, Lynn Margulis (the world's authority on endosymbiosis) is also my idea of a real scientist. She absolutely loves getting knee-deep in the mud of some tropical lagoon and scraping scum off of rocks to look at under the microscope. She's spent her entire professional life studying microorganisms in the field and in the lab, and has revolutionized our understanding of the evolutionary biology of microorganisms. Like Harry, she's the epitome of an evolutionary biologist, because she also "a feel for the organism" which leads her to discover things nobody ever thought to look for before, such as symbiotic bacteria embedded in the cell membranes of symbiotic protozoa from the guts of termites.

I have yet to meet or hear about or read about any ID supporter who does anything like what Harry and Lynn do. Yes, Michael Behe is a biochemist, but the things he does in his laboratory at Lehigh have little or nothing to do with ID. And William Dembski wouldn't know an actual living organism if it lunged out and bit him on the ankle.

Biology, and especially evolutionary biology, is that branch of the natural sciences founded and maintained by people who loved and were obsessed with nature and natural things. Darwin and Wallace and Fisher and Haldane and Wright and Dobzhansky and Mayr and Simpson and Stebbins and Hamilton and Trivers and Margulis and the two Wilsons (Edward O. and David Sloan): these are my heroes, and they are the "naturalists" (see how the word has another, much more positive meaning?) who have been the inspiration for my research, insignificant as it is compared with theirs.

And all that IDers can generally do is say "No, you're wrong, it can't happen that way, in fact it can't happen at all without a deus ex machina?" Ugh: boring, pointless, and most of all, no "feel for the organism".

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As always, comments, criticisms, and suggestions are warmly welcomed!

--Allen

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