EVOLUTION AND THE ID WADS

Endless Forms Most Beautiful

A New Revolution in Biology

Over the past two decades, a new revolution has unfolded in biology. Advances in embryology and
evolutionary development biology (dubbed Evo Devo)—involving genetic switches and simple rules
that shape animal form and evolution—have profoundly reshaped our picture of how evolution works.

SEAN B. CARROLL

An excerpt from
Endless Forms Most Beautiful: The New Science ofEvo Devo.

T
he great variety in the size, shape, organization, and
color of animal bodies raises deep questions about
the origins of animal forms. How are individual
forms generated? And, how have such diverse forms evolved?
These are very old questions in biology that date back to
Darwin's time and before, but only very recently have deep
answers been discovered, many of them so surprising and
profound that they have revolutionized our views on the
making of the animal world and our place in it. We all share
an attraction to animal form, but my aim is to expand that

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wonder and fascination to how form is created, that is, was studying genetic variation in populations, igno-
to our new understanding of the biological processes rant of the relationship between genes and form.
that generate pattern and diversity in animal design. Perhaps even worse, the perception of evolutionary
While Darwin, Huxley, and their allies knew that biology in some circles was that it had become rele-
embryological development was key to evolution, for gated to dusty museums.
more than one hundred years after their chief works, Such was the setting in the 1970s when voices for
virtually no progress was made in understanding the the reunion of embryology and evolutionary biology
mysteries of development. The puzzle of how a simple made themselves heard. Most notable was Stephen
egg gives rise to a complete individual stood as one of Jay Gould, whose book Ontogeny and Phylogeny
the most elusive questions in all of biology. Many revived discussion of the ways in which the modification of
thought that development was hopelessly complex, and development may influence evolution. Gould had also stirred up
would involve entirely different explanations for different types of evolutionary biology when, with Niles Eldredge, he took a fresh
animals. So frustrating was the enterprise, that the study of look at the patterns of the fossil record and forwarded the idea of
embryology, heredity, and evolution, once intertwined at the core "punctuated equilibrium"—that evolution was marked by long
of biological thought a century ago, fractured into separate fields periods of stasis (equilibrium) interrupted by brief intervals of
as each sought to define its own principles. rapid change (punctuation). Gould's book and his many subse-
quent writings reexamined the "Big Picture" in evolutionary biol-
Because embryology was stalled for so long, it played no part
ogy and underscored the major questions that remained unsolved.
in the so-called "Modern Synthesis" of evolutionary tiiought that
He planted seeds in more than a few impressionable young scien-
emerged in the 1930s and 1940s. In the decades after Darwin,
tists, myself included.
biologists struggled to understand the mechanisms of evolution.
At the time of The Origin of Species (1859), the mechanism for the To me, and others who had been weaned on the emerging suc-
inheritance of traits was not known. Mendel's work was redis- cesses of molecular biology in explaining how genes work, the sit-
covered decades later and genetics did not prosper until well into uations in embryology and in evolutionary biology were both
the 1900s. Different kinds of biologists were approaching evolu- unsatisfying states of affairs, and enormous potential opportuni-
tion at dramatically different scales. Paleontology focused on the ties. Our lack of embryological knowledge seemed to make much
largest time scales, the fossil record, and the evolution of higher of the discussion in evolutionary biology about the evolution of
taxa. Systematists were concerned with the nature of species and form just futile exercises in speculation. How could we make
the process of speciation. Geneticists generally studied variation in progress on questions involving the evolution of form without a
traits in just a few species. These disciplines were disconnected scientific understanding of how form is generated in the first
and sometimes hostile over which offered the most worthwhile place? Population genetics had succeeded in establishing the prin-
ciple that evolution is due to changes in genes, but this was a prin-
insights into evolutionary biology. Harmony was gradually
ciple without an example. No gene that affected the form and
approached through an integration of evolutionary viewpoints at
evolution of any animal had been characterized. New insights in
different levels. Julian Huxley's book Evolution: The Modern
evolution would require breakthroughs in embryology.
Synthesis (1942) signaled this union and the general acceptance of
two main ideas. First, that gradual evolution can be explained by
small genetic changes that produce variation which is acted upon The Evo-Devo Revolution
by natural selection. And second, that evolution at higher taxo- Everyone knew that genes must be at the center of the myster-
nomic levels and of greater magnitude can be explained by these ies of both development and evolution. Zebras look like zebras,
same gradual evolutionary processes sustained over larger periods. butterflies look like butterflies, and we look like we do because
The Modern Synthesis established much of the foundation for of the genes we carry. The problem was that there were no, or
how evolutionary biology has been discussed and taught for the very few, clues as to which genes mattered for the development
past sixty years. However, despite the monikers of "Modern" and of any animal.
"Synthesis," it was incomplete. At that time, we could say that The long drought in embryology was eventually broken by a
forms do change, and that natural selection is a force, but we few brilliant geneticists who, while working with the fruit fly, the
could say nothing about how forms change, about the visible
drama of evolution as depicted, for example, in the fossil record. Sean B. Carroll is an investigator at the Howard Hughes Medical
The Synthesis treated embryology as a "black box" that somehow Institute and a professor of genetics at the University of
transformed genetic information into three-dimensional, func- Wisconsin-Madison. He is one of the leading biologists of his gen-
tional animals. eration, and his seminal discoveries have been featured in many
The stalemate continued for several decades. Embryology was popular publications. This article is excerpted by permission from
preoccupied with phenomena that could be studied by manipu- his new book Endless Forms Most Beautiful: The New Science
lating the eggs and embryos of a few species, and the evolutionary of Evo Devo (WW Norton, New York, 2005). E-mail sbcarroll@
framework faded from embryology's view. Evolutionary biology wisc.edu.

SKEPTICAL INQUIRER November / December 200S 49

 EVOLUTION AND THE ID WARS

workhorse of genetics for the past eighty years, devised schemes to aspects of fruit fly body organization were found to have exaa
find die genes that controlled fly development. The discovery of counterparts that did die same thing in most animals, including
these genes and their study in the 1980s gave birth to an exciting ourselves. This discovery was followed by the revelation mat die
new vista on development, and revealed a logic and order under- development of various body parts such as eyes, limbs, and hearts,
lying the generation of animal form. vastly different in structure among animals and long thought to
Almost immediately after the first sets of fruit fly genes were have evolved in entirely different ways, were also governed by the
characterized came a bombshell that triggered a new revolution in same genes in different animals. The comparison of developmen-
evolutionary biology. For more than a century, biologists had tal genes between species became a new discipline at the interface
assumed that different types of animals were genetically con- of embryology and evolutionary biology—evolutionary develop-
structed in completely different ways. The greater the disparity in mental biology or "Evo-Devo" for short.
animal form, die less (if anything) the development of two ani- The first shots in the Evo-Devo revolution revealed diat
mals would have in common at the level of their genes. One of despite their great differences in appearance and physiology, all
the architects of the Modern Synthesis, Ernst Mayr, had written complex animals—flies and fly catchers, dinosaurs and trilobites,
diat "the search for homologous [the same] genes is quite (utile butterflies and zebras and humans—all share a common "toolkit"
except in very close relatives." But contrary to the expectations of of "master" genes that govern the formation and patterning of
any biologist, most of the genes first identified as governing major their bodies and body parts. The important point to appreciate is
that its discovery shattered our previous notions of animal rela-
tionships and of what made animals different, and opened up a
whole new way of looking at evolution.
Evo Devo: We now know from sequencing the entire DNA of species
A Revolutionary Quartet (their "genomes") that not only do flies and humans share a large
cohort of developmental genes, but that mice and humans have
The impact and importance of Evo Devo emerges virtually identical sets of about 25,000 genes, and diat chimps and
from four points: humans are nearly 99 percent identical at the DNA level. These
First, Evo Devo constitutes a third major act in a facts and figures should be humbling to diosc who wish to hold
continuing evolutionary synthesis. Evo Devo has humans above die animal world and not an evolved part of it. I
not just provided a critical missing piece of the wish the view I heard expressed by Lewis Black, the stand-up
"Modern Synthesis"—embryology—and integrated
comedian, was more widely shared. He said he won't even debate
it with molecular genetics and traditional elements
such as paleontology. The wholly unexpected evolutions detractors because "We've got die fossils. Wc win." Well
nature of some of its key discoveries and the put, Mr. Black, but there is far more to rely on than just fossils.
unprecedented quality and depth of evidence it has Indeed, die new facts and insights from embryology and Evo-
provided towards settling previously unresolved Devo devastate lingering remnants of stale anti-evolution rhetoric
questions bestow it with a revolutionary character. about the utility of intermediate forms or the probability of evolv-
Second, Evo Devo provides a new means of teach- ing complex structures. We now understand how complexity is
ing evolutionary principles in a more effective frame- constructed from a single cell into a whole animal. And we can
work. By focusing on the drama of the evolution of see, widi an entirely new set of powerful methods, how modifica-
form, and illustrating how changes in development tions of development increase complexity and expand diversity.
and genes are the basis of evolution, the deep princi-
The discovery of the ancient genetic toolkit is irrefutable evidence
ples underlying the unity and diversity of life emerge.
Furthermore, the visible forms of gene expression of die descent and modification of animals, including humans,
patterns in embryos and the concrete inventories of from a simple common ancestor. Evo-Devo can trace the modifi-
toolkit gene sets in different species provide more cations of structures through vast periods of evolutionary time—
effective ways of illustrating evolutionary concepts to see how fish fins were modified into limbs in terrestrial verte-
than previous, more abstract approaches. brates, how successive rounds of innovation and modification
crafted mouthparts, poison claws, swimming and feeding
Third, because Evo Devo reveals and illustrates
the evolutionary process and principles in such tan- appendages, gills, and wings from a simple tube-like walking leg,
gible ways, it has a critical role to play in the fore- and how many kinds of eyes have been constructed beginning
front of the societal struggle over the teaching of with a collection of photosensitive cells. The wealth of new data
evolutionary biology. from Evo-Devo paints a vivid picture of how animal forms are
And finally, the importance of evolutionary biol- made and evolve.
ogy is far more than mere philosophy. The fate of
the endless forms of nature, including humans,
depends on a broader understanding of humans' The Toolkit Paradox and the Origins of Diversity
impacts on evolution. The stories of shared body-building genes and of the similarities
of our genome to that of other animals have slowly been gaining

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 EVOLUTION AND THE ID WARS

in public awareness. What is gen- ulatory. This DNA determines

erally neglected, however, is how when, where, and how much of
the discovery of this common a gene's product is made.
toolkit and of great similarities Regulatory DNA is organized
among different species' genomes into fantastic little devices that
presents an apparent paradox. If integrate information about
the sets of genes are so widely position in the embryo and the
shared, how do differences arise? time of development. The out-
The resolution of this paradox put of these devices is ultimately
and its implications are central to transformed into pieces of
anatomy diat make up animal
my story. The paradox of great
forms. This regulatory DNA
genetic similarity among diverse
contains the instructions for
species is resolved by two key
building anatomy, and evolu-
ideas. These concepts are crucial
tionary changes within this reg-
for understanding how die
ulatory DNA lead to die diver-
species-specific instructions for sity of form.
building an animal are encoded
in its DNA and how form is gen-
erated and evolves. The substance The Grandeur in a More
of these ideas has received scant, Modern Synthesis: Act III
if any, attention in the general The continuing story of evolu-
press, but they have profound tion may be thought of as a
implications for understanding drama in at least diree acts. In
great episodes in life's history Act I, almost 150 years ago,
such as the explosion of animal Darwin closed the most impor-
forms during the Cambrian tant book in the history of biol-
period, the evolution of diversity ogy by urging his readers to see
within groups such as butterflies die grandeur in his new vision of
or beetles or finches, and our evolution from a common ancestor nature [see box]. In Act II, the architects of the Modern Synthesis
with chimps and gorillas. unified at least diree disciplines to forge a grand synthesis. Here in
The first idea is that diversity is not so much a matter of the Act III, there is also a special grandeur in die view embryology and
complement of genes in an animal's toolkit, but in the words of evolutionary developmental biology provide into the making of
Eric Clapton, "It's in the way that you use it." The development animal form and diversity. Pan of it is visual, in that we can now
of form depends upon the turning on and off of genes at differ- see how die forms of different animals actually take shape.
ent times and places in die course of development. Differences in But beauty, in science, is much more than skin deep. The best
form arise from evolutionary changes in where and when genes science is an integrated product of our emotional and intellectual
are used, especially those genes that affect the number, shape, or sides, a synthesis between what is often referred to as our "left"
size of a structure. There are many ways to change how genes are brain (reasoning) and "right" brain (emotional/artistic) hemi-
used and diat this has created tremendous variety in body designs spheres. The greatest "eurekas" in science combine both sensual
and die patterning of individual structures. aesthetics and conceptual insight. The physicist Victor Weisskopf
The second idea concerns where in the genome the "smoking noted, "What is beautiful in science is the same diing that is beau-
guns" for die evolution in form are found. It turns out that it is tiful in Beethoven. There's a fog of events and suddenly you see a
not where we have been spending most of our time for die past connection. It expresses a complex of human concerns that goes
forty years. It has long been understood that genes are made up of deeply to you, diat connects things that were always in you that
long stretches of DNA diat are decoded by a universal process to were never put together before."
produce proteins, which do the actual work in animal cells and Those connections are revealing some simple, elegant truths
bodies. The genetic code for proteins, a twenty-word vocabulary, that deepen our understanding of all animal forms, including our-
has been known for forty years and it is easy for us to decode selves, and should transform how evolutionary science is taught
DNA sequences into protein sequences. What is much less appre- and accepted.
ciated is that only a tiny fraction of our DNA, just about 1.5 per-
cent, codes for the roughly 25,000 proteins in our bodies. So what Evo Devo and the Teaching of Evolution
else is diere in die vast amount of our DNA? Around 3 percent of The teaching of evolution faces two challenges. The first is diat it
it, made up of about one hundred million individual bits, is reg- is a vast and growing subject that encompasses many disciplines.

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 EVOLUTION AND THE ID WADS

The second is that it is actively opposed, particularly in die U.S., played a central role in the development of all evolutionary
by some (but not all!) religious factions. I will address die new diought. The advantages of an embryological approach to teach-
contribution Evo Devo can make to improving general public ing evolution are several-fold.
understanding first, die issue of opposition later. First, it is a small leap to go from the building of complexity
In general, die public understanding of evolution in the in one generation from egg to adult to appreciating how incre-
United States is particularly abominable. In a survey of citizens in ments of change in the process, assimilated over greater time peri-
twenty-one countries regarding general environmental and scien- ods, produce increasingly diverse forms.
tific knowledge, die U.S. placed dead last on die question of Second, we now have a very firm grasp of how development is
human evolution. Looking at die bright side, the U.S.A. can only controlled. We can explain how toolkit proteins shape form, that
move up from here. toolkit genes are shared by all animals, and diat differences in
In another survey, by die National Science Board in 1996, 52 form arise from changing the way diey are used. The principle of
percent of Americans polled either agreed with (32 percent) or did descent by modification (of development) is crystal clear.
not know (20 percent ) whedier die statement "The earliest Third, an enormous practical advantage is the visual nature of
humans lived at die same time as the dinosaurs" was true or not. the Evo Devo perspective. The Chinese proverb that "hearing
Score diat fact as two points for The Flintstones, zero for about something a hundred times is not as good as seeing it once"
Darwin, Huxley, and the educational system of the world's most is sound educational doctrine. We learn more by combining visu-
wealthy, powerful, and technologically driven nation. als widi text. Let's show students embryos and how stripes, spots,
The scandal of diis ignorance is on par, I would say, with not and all die glorious features of animal forms are made. The evo-
knowing how the U.S.A. was formed, the content of its lutionary concepts follow naturally.
Constitution, or the roots of Western civilization. This knowledge A fourth benefit of diis approach is diat it brings genetics
is considered basic literacy and taught and repeated at many grade much closer to the powerful evidence of paleontology. Dinosaurs
levels. So, too, is biology and earth science for which evolution and trilobites are the poster children of evolution diat inspire the
must provide the basic framework. Yet, die statistics are appalling. vast majority of diose who touch diem. By placing these wonders
The situation is bad enough, and reflected in other figures of die ancient past in a continuum from die Cambrian to die pre-
about science and math literacy, that the blame can probably be sent life's, history is made much more tangible. It would indeed
shared in many quarters. There are plenty of books written about be a wonderful world if every student had guided, repeated class-
and organizations studying the general problem of scientific illit- room contact with some fossils.
eracy and its causes. I won't get into finger pointing here. The only Let me offer a couple more general suggestions. Natural selec-
way up is through education. I would rather focus on what biol- tion is often at best described as a "just-so story" of adaptations.
ogists and their allies at all levels of the teaching profession can do Finches beaks changed due to die type of food available, modis got
to improve matters, particularly in regard to evolution. darker because of pollution, etc. But I do not diink diat die power
First, we must insist that evolution is much more than just a of small increments of selection, compounded over hundreds or
topic in biology, it is die foundation of the entire discipline. thousands of generations, is widely taught or understood. The
Biology without evolution is like physics widiout gravity. Just as commonly repeated phrase "survival of die fittest" connotes more
we cannot explain the structure of the universe, die orbits of die of a gladiator contest dian the subde power of selection to act on
planets and moon, or tides from mere measurement, we cannot very small differences in overall survival and fecundity. The spread
explain human biology or Earths biodiversity via a compendium of favorable mutations in populations is easily simulated and illus-
of diousands of little facts. All general survey courses and texts trated, and it underscores die time dimension of evolution.
must have evolution as their central unifying theme. Finally, at die university level, the evolutionary view of life
With respect to die scientific content to be taught, Evo Devo should be as fundamental to a college degree as Psychology 101 or
has much to contribute that is new, tangible, and convincing. Western Civilization. But rather than asking students to memo-
Since die Modern Synthesis, most expositions of die evolutionary rize and regurgitate mountains of testable facts, we should empha-
process have focused on microevolutionary mechanisms. Millions size study of the history of the discovery of evolution, its major
of biology students have been taught the view (from population characters and ideas, and the basic lines of evidence. This would
genetics) that "evolution is change in gene frequencies." Isn't that do far more to inform citizens and prepare teachers than forcing
an inspiring theme? This view forces the explanation toward students to remember the Latin names of species. We are stoning
mathematics and abstract descriptions of genes, and away from our children to utter boredom widi little pebbles and missing the
butterflies and zebras, or Australopithecines and Neanderthals. big picture. The drama of the story of evolution will recapture stu-
The evolution of form is the main drama of life's story, both as dent interest.
found in the fossil record and in die diversity of living species. So, There is, especially in the U.S.A, another obstacle besides con-
let's teach that story. Instead of "change in gene frequencies," let's tent and teaching mediods to evolutionary literacy. I will address
try "evolution of form is change in development." This is, of diat next. But, even widiout the active opposition, we can do bet-
course, a dirowback to the Darwin-Huxley era, when embryology ter, and we have to do better.

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 EVOLUTION AND THE ID WADS

Evo D e v o a n d t h e Evolution/Creation Struggle Box: The Biochemical Challenge to Evolution. As a credentialed sci-
In the short time between the first and second edition of The entist, Behe's book was received as a godsend by creationists and
Origin of Species, Darwin inserted three more words into that is perhaps the best known treatment of Intelligent Design. But
famous closing paragraph, adding "by the Creator" to rewrite the Behe's main claim, that the living cell is an entity of irreducible
phrase as "having been originally breathed by the Creator into a complexity, is empty. Behe was counting on biology to hit a wall
few forms or into one . . . ." Darwin later expressed his regret for in reducing complex phenomena to molecular processes. H e joins
doing so in a letter to botanist J . D . Hooker: a long line o f prognosticators whose pessimistic forecasts have
been obliterated in the continuing revolution in the life sciences.
But I have long regretted that I truckled to public opinion, and
Scott Gilbert, a biologist at Swarthmore College, author of the
used the Pentateuchal term of creation, by which I really meant
"appeared" by some wholly unknown process. leading college text of developmental biology, and accomplished
historian of embryology and evolutionary biology, has summa-
The insertion of these words was intended to appease critics rized the Behe position, and its failure as follows:
and make Darwin's evolutionary ideas more palatable. It has cer-
To creationists, the synthesis of evolution and generics cannot
tainly served to fuel much speculation about Darwin's actual reli-
explain how some fish became amphibians, how some reptiles
gious views. For some, this olive branch and Darwin's reticence in became mammals, or how some apes became human.. . . Behe
disclosing his beliefs (which are only revealed to some degree in pri- named this inability to explain the creation of new taxa through
vate correspondence and unpublished notebooks) were the foun- genetics "Darwin's black box." When the box is opened, he
expects evidence of the Deity to be found. However, inside
dation for reconciling and accommodating evolution and religion.
Darwin's black box resides merely another type of genetics—
Plenty of scientists and a broad spectrum of religious denom- developmental genetics.
inations have found such an accommodation. For example, in
1996, Pope John Paul II reiterated the Catholic position that the Developmental genetics has been shedding new light on the
h u m a n body has evolved according to natural processes. But making of complexity and the evolution of diversity for twenty
while some denominations have explicitly accepted the reality of years. Creationists just plain refuse to see it. H o w is such overt evi-
biological evolution, fundamentalists who insist upon a literal dence ignored or dismissed? I can't pretend to understand the psy-
reading of the Bible (referred to as "Creationists") and propo- chological mechanisms that allow humans to deny reality. But I
nents of "stealth" Creationist ideas such as Intelligent Design do understand the desperate political and rhetorical tactics of
remain firmly opposed to evolutionary science and actively pro- those who, holding a losing hand, refuse to accept it.
mote legislation aimed at crippling the teaching of evolution in As exasperating as the continuous battle with creationists may
public schools. seem, the scientific community is now better organized and more
Goethe said that "Nothing is worse than active ignorance," prepared to deal with the movement. But the batde against igno-
and it is the agenda of these lost souls that the scientific and edu- rance is not won. Q
cational communities must thwart. I want to be very clear here in
my position. I believe that the teaching of evolution and science
are best served by promoting the scientific method and scientific
knowledge and not by attacking religious views. T h e latter is a
A Grandeur . . . and Endless
futile, counterproductive battle. However, I also believe, as many Forms Most Beautiful
denominations have also concluded, that religion is better served
by promoting and evolving their respective teachings and theolo- Darwin closed t h e first e d i t i o n o f The Origin of
Species w i t h w h a t has become perhaps t h e most
gies, and not by attacking science, which is definitely a losing
w i d e l y q u o t e d passage in all o f biology:
strategy.
Charles Harper, executive director of the John Templeton There is a grandeur in this view of life, with its sev-
eral powers, having been originally breathed into a
Foundation, an organization interested in the relationship few forms or into one: and that whilst this planet
between theology and science, wrote recendy in the leading sci- has gone cycling on according to the fixed law of
gravity, from so simple a beginning endless forms
ence journal Nature that:
most beautiful and most wonderful have been, and
As scientific knowledge grows, religious commitments predicated are being, evolved.
on "gaps" in scientific understanding will invariably shrink as
those gaps arc closed. Those Christians who are currendy fighting I have chosen f o u r words t h a t remained com-
evolutionary science will eventually need to take it seriously. pletely u n t o u c h e d t h r o u g h o u t all versions a n d edi-
tions, "endless forms most b e a u t i f u l , " as t h e inspi-
Harper is right. In this time of unprecedented power in under- r a t i o n f o r and t h e m e o f this article. This phrase res-
standing embryos, genes, and genomes, and with the continual onates most perfectly w i t h t h e t h e m e of my book
expansion of the fossil record, those gaps are fast disappearing. and captures t h e essence o f t h e n e w science o f Evo
O n e example of a mistaken faith in those gaps is dial of bio- Devo.
chemist Michael Behe who in 1996 published Darwin's Black

SKEPTICAL INQUIRER November / December 200S S 3