FOR E WOR D

Becoming is superior to being.

—Paul Klee

M etamorphoses have long held a spell over the human imagi-

nation. For the ancients, as chronicled by the Roman writer
Ovid in his Metamorphoses, the gods could assume any and many
forms, like the comic-strip superhero Metamorpho, able to shape-
shift his entire being into a mere part of the human body. Drowned
Narcissus is turned into a yellow-and-white flower, the narcissus.
Phaethon’s sisters are metamorphosed into poplar trees, their tears
hardened into drops of amber. Jupiter transforms the nymph Io into
a heifer to conceal her. Daphne eludes Apollo by changing into a
laurel tree, the lovelorn god winding up with laurel leaves in his hair.
Neptune turns Cygnus into a swan—and so it goes, one organism
transforming into another, pushed by desire rather than regard for
academically immutable species lines. In fact, one might argue that
Aristotle, often called the first biologist, rejected natural selection
because he associated it with the unsavory superstitions of his imagi-
native Greek forebears, especially Empedocles, whose early version
of natural selection featured “man-faced ox-progeny” that must have
reminded Aristotle more of myth than science. The fifth-century B.C.
philosopher Empedocles of Acragas (now Sicily) devised a theory in
which organs, arising outside bodies, first roamed the land, the less
fit ones dying out and the remaining ones coalescing to make bodies.
Charles Darwin knew about Empedocles’ theory and its rejection
by Aristotle, whose writings Darwin’s friend translated so that he
could print it in The Origin of Species to show the cluelessness of
the ancients.
In keeping with Aristotle’s throwing out the baby of natural selec-
tion with the bathwater of mythical hybrid beings, metamorphosis—

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 xiv  The Mystery of Metamorphosis

from the Greek words meta (μετα), “after,” and morphe (μορφή ),
meaning “form”—continues, for the most part, to be considered
fantasy in popular culture. Except for some details of insect biol-
ogy, it is considered a kind of magic or reincarnation, a subject for
fiction not science. In Kafka’s The Metamorphosis, perhaps the
most famous short work of fiction ever written, Czech clerk Gregor
Samsa wakes up transformed into a giant cockroach; he is unable
to roll over but still conscious of his human family, to whom he
listens in the next room. Britain’s Prince Philip is alleged to have
stated that he wished to reincarnate as a deadly virus in order to
reduce the world’s population; and, in a television-age throwback to
the strange comeuppances doled out by the gods, the womanizing
character Quagmire, in the cartoon Family Guy, is reincarnated as
a prophylactic.
However, metamorphoses are real. The striking metamorphoses
of organisms are not confined to butterflies and children’s story-
books, but apply to sea creatures, many species of insects, and argu-
ably, in a muted or “incomplete” form, to us when we undergo the
difficult, hormonally mediated transition from preteen to adult. Like
Aristotle, Frank Ryan, an evolutionary biologist as well as a medical
doctor, dispenses with the gods—but not with natural selection—as
he explicates the “beautiful mystery” of nature’s transformations.
And he resuscitates the seemingly primitive idea that Aristotle
dismissed, of radical hybrids.
Ryan discusses the personalities, by turns bold and retiring, social
and introverted, confrontational and easygoing, of key figures in our
emerging understanding of the chemical—and evolutionary—bases
for metamorphoses. Ryan globe-trots to meet the key figures in this
biological subfield who’ve piqued his—and through him, our—
curiosity. When they are not alive, he talks to their students and
family members; and an expert himself, a doctor who has pioneered
new evolutionary concepts about viruses and written a bestseller
on tuberculosis, Ryan reads the primary scientific literature before
relaying the amazing findings and open questions to us. Here we
encounter the greats of classical metamorphoses studies, from the

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 Foreword   xv

observations of Jean-Henri Fabre, the nineteenth-century author of

The Life of the Caterpillar to the appropriately named Vincent B.
Wigglesworth, who pioneered our understanding of hormonal
transformation in insects, the chemical essence of insect pubes-
cence. At first glance a little-known story such as this one might
seem of interest only to a small coterie of specialist academics. But
Ryan shows that the extent of the research of Wigglesworth and
his students reaches far and wide—into the manufactured paper of
the New York Times, Wall Street Journal, Boston Globe, Science,
and Scientific American, for example, which was shown to contain
plant-produced compounds that stop fire bug larvae from undergo-
ing metamorphosis.
The case of the book’s other major protagonist, Donald Williamson,
is wilder. For Williamson has offered a new theory of metamorpho-
sis that to mainstream biology is as easy to dismiss as Aristotle was
for Darwin—or Empedocles for Aristotle. Williamson iconoclasti-
cally proposes metamorphosis comes not of the “gradual accumula-
tion of mutations” over millions of years, as neo-Darwinism would
have it, but issues rather from “forbidden fertilizations”—the joining
of genomes of completely different species in new, sometimes viably
reproducing bodies.
Williamson’s radical proposed explanation sounds less fantas-
tic if you realize that many nonmammalian animals fertilize eggs
outside their bodies. For example, it is completely within the realm
of possibility that flying insects in the late Paleozoic swampy trop-
ics dropped some of their sperm on the egg masses of velvet worms.
Velvet worms, called Onychophora, abound in tropical leaf litter
on the forest floor. They extrude gummy, sticky stuff to help them
catch edible prey. Velvet worms are tropical, crawling, many-legged
beings who would look like caterpillars to your mother. Most of the
time, such inappropriate sperm drops by an oversexed impatient flut-
terer would have had no effects, but not always. Sometimes sperm
and egg unions of very distantly related animals are neither infertile
nor aborted. Williamson was impressed by the fact that juvenile
forms of very different species can be amazingly similar given the

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 xvi  The Mystery of Metamorphosis

extreme differences of the adults they become. Following his curios-

ity, he tried to interbreed different species of marine invertebrates:
he fertilized eggs of an ascidian (Ascidia mentula) with sperm of
a sea urchin (Echinus esculentus). In one experiment all the eggs
developed as easel-shaped pluteus larvae—sea urchin larvae from
ascidian eggs. About 8 percent of these plutei grew into sea urchins,
but the other 92 percent resorbed their arms and became spher-
oids, each with a sucker. Why does this matter? Because no adult
or larval sea urchin ever grows any head suckers—but the tadpole
larvae of ascidians do. (Later experiments crossing the sea urchin
Psammechinus miliaris with the heart urchins Echinocardium
cordatum were recorded in the documentary Hopeful Monsters
by Robert Sternberg of Imperial College, University of London.)
In Williamson’s view, the overlapping genomes express themselves
with a time lag in development. The pluteus, tadpole, caterpillar,
or other animal genes of the larval form are expressed first; then,
exquisitely sensitive to environmental change, they shut off, and the
adult gene expression begins. This is metamorphosis.
If accepted, Williamson’s conclusions will require reorganiza-
tion of the entire science of zoology. The most telling proof that
Williamson is correct lies in the appearance of head suckers (“adhe-
sive papillae”) in two sets of his hybrids. Head suckers typically
appear (in ascidians) during metamorphosis when fishlike swim-
ming larvae transform into sessile bottom-dwelling adults. But now
why would fish-tailed larvae, which resemble our vertebrate ances-
tors, transform into decidedly un-fishlike adults that produce eggs
and sperm but, attached to rocks, never travel again? The situation
is one of those tellingly imperfect would-be designs, like the wings
of flightless birds, that clue us in to the fact that the organism before
us is the result of peculiar and unexpected history.
Williamson’s work suggests that the adult forms of metamorphos-
ing species—the radially symmetrical sea urchin on the sea bottom,
the frog leaping about the bank of a forest pond, the flighty butterfly
fluttering and hovering in the air—are literally alien genomes with
respect to the larval forms. Interspecies beings, these organisms

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 Foreword   xvii

go through such profound changes because they have successfully

merged stranger genomes. Such Prince Charming transformations
seem to testify to quite complicated ancient relationships; they are
emblems of the most boggling sort of “mixed marriages” that,
despite their starkly different backgrounds, somehow genetically
worked it out.
Whether or not Williamson’s ideas (wholly or in part) ultimately
become accepted, the fact remains that, unlike the fairy tale in
which the princess kisses the frog, these larval forms really do meta-
morphose. Readers of Ryan’s book will find that these metamor-
phoses remain a mystery—but less so than when they started the
book. Ryan makes the beauty of metamorphosis palpable, and lays
down the work of those variant personalities—by turns taciturn
and gregarious, methodical and intuitive—who’ve attempted to
wrest from nature the secrets of ontogenetic transformation. Apart
from the practical, philosophical, and evolutionary ideas broached
and explored, Ryan’s work is a model of informed scientific history.
He has the advantage of being well versed in the difficult terrain
that he traverses. But he is also able to write clearly, to synthesize
broadly, while still resisting the temptation to foreclose or come
to too-precipitous conclusions. In an intricate but always fascinat-
ing and detailed account of metamorphoses and the scientists who
study them, Ryan presents both an ethnography and a history, not
only of science as it is believed and accepted to have happened after
the fact, but also of science in medias res—science right now, before
our eyes, forming—emerging and groping toward new understand-
ings as if it were itself some sort of metamorphosing being, rustling
in its pupa before emerging resplendently into the adulthood and
newfound symmetry of a deeper understanding of biological process
and the natural histories that mark it.

— Dorion Sagan and Lynn Margulis

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