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The Fantastic Laboratory of Dr Weigl How Two Brave
Scientists Battled Typhus and Sabotaged the Nazis 1st
Edition Arthur Allen Digital Instant Download
Author(s): Arthur Allen
ISBN(s): 9780393081015, 039308101X
Edition: 1
File Details: PDF, 3.64 MB
Year: 2014
Language: english
 THE
FANTASTIC
LABORATORY
OF
DR.
WEIGL

HOW TWO BRAVE SCIENTISTS BATTLED TYPHUS

AND SABOTAGED THE NAZIS

ARTHUR ALLEN

W. W. NORTON & COMPANY

New York | London

2
To Margaret, Ike, and Lucy

3
 CONTENTS

Preface

Introduction

Chapter One: Lice/War/Typhus/Madness

Chapter Two: City on the Edge of Time

Chapter Three: The Louse Feeders

Chapter Four: The Nazi Doctors and the Shape of Things to Come

Chapter Five: War and Epidemics

Chapter Six: Parasites

Chapter Seven: The Fantastic Laboratory of Dr. Wiegl

Chapter Eight: Armies of Winter

Chapter Nine: The Terrifying Clinic of Dr. Ding

Chapter Ten: “Paradise” at Auschwitz

Chapter Eleven: Buchenwald: Rabbit Stew and Fake Vaccine

Chapter Twelve: Imperfect Justice

Afterword

Notes

Acknowledgments

Index

4
5
6
 THE
FANTASTIC
LABORATORY
OF
DR. WEIGL

7
 PREFACE

A few years ago I found myself in a dim corridor at the Institute of

Epidemiology and Hygiene in Lviv, Ukraine, trying to persuade Dr.
Oleksandra Tarasyuk, the institute’s polite but recalcitrant director, to let
me watch the feeding of the lice.
Why, one might ask, would anyone come all the way to Ukraine to
look at lice? They are, after all, common loathsome insects, synonymous
everywhere with disease and filth, wretchedness and neglect. To the
naked eye, Dr. Tarasyuk’s lice were no different from the ones that I and
millions of other parents combed from the scalps of our children during
elementary school infestations. I had once or twice examined my
children’s cohabitants under a microscope and found them to be
surprisingly intricate, greasy-brown creatures whose guts contained tiny
but distinct canals of blood.
But the lice of 12 Zelena Street, Lviv, were not quite ordinary
creatures. For one thing, they were body lice (Pediculus humanus
humanus) rather than head lice (Pediculus humanus capitis) . The two
insects, varieties of a single species, are remarkably similar; even
geneticists have trouble parsing their essential difference. Both nourish
themselves by poking an exoskeletal needle into the warm skins of
humans and employing the musculature of their tiny proboscides to
extract our blood. But for reasons that biologists have yet to understand,
there is one fundamental distinction between head lice and body lice:
head lice are a nuisance, but only body lice transmit one of humankind’s
most fearsome diseases, typhus. Body lice are thus players, in a way that
head lice have never been, in some of the great tragedies and most
horrendous pages of history. Few organisms have been as deadly to
doctors and medical researchers as typhus. This is perhaps not surprising,
because the sick shed lice, which are fussy about heat and cold and
abandon the body once temperatures fall below 98 degrees Fahrenheit or
rise above 102 degrees, desperately searching for a new home. Each
laboratory in the fight against typhus had its martyrs, and publications
about the disease were inevitably dedicated to fallen colleagues. This
explains why the lice of Lviv led such a charmed existence, spending most
of their lives swarming together in the comfort of heated wooden

8
cabinets, unlike the rootless lumpen proletarians that burrow, for a brief
while, in the hair of schoolchildren.
Like many scientists who spend careers in close proximity to lab
animals, Dr. Tarasyuk felt quite protective of hers. “It’s very difficult to
keep this population alive,” she told me in a remorseful tone. “ They need
particular temperature levels at different stages of their lives. They feed
only once a day. And we have to make sure that the feeders are healthy.”
And what do these lice eat? Human blood, of course. And how do they
procure it? Why, by being placed in cages on the legs of human beings.
The feeders at the institute, each paid a small sum for their sacrifice and
blood donation, were mostly lab technicians. The idea of letting me, a
stranger who didn’t even speak Ukrainian, into the lab filled Dr. Tarasyuk
with horror. I could give the lice a disease! I could threaten the survival of
the colony! She could lose her job! “Come back the next time you are in
Lviv, but give us more warning,” she told me with a frown. “We would
love to see you again.”
Afterwards I stood outside the rather plain, five-story, Bauhaus-
influenced building for a few moments and tried to conjure up a picture of
its past. The lice of 12 Zelena Street are the descendants of a colony bred
seven decades ago by the zoologist Rudolf Weigl, who crossed lice picked
from the bodies of Russian prisoners of World War I with those nestled in
the robes of Ethiopian highlanders. With these lice and a lot of ingenuity,
Weigl in the 1920s created the first effective vaccine against typhus, a
disease that terrorized the world, inspired the creation of Zyklon B gas,
and provided a pretext for the worst human crimes in history. Weigl’s
discovery drew global notice. Nobel laureates trod the corridors of his
institute to study his techniques and pay homage to him. Agents from the
Nazi SS and the Soviet NKVD sniffed around the halls of his institute;
Nikita Khrushchev, later the Soviet premier, and Hans Frank, the Nazi
governor of Poland, appeared at his doors, soliciting Weigl’s services.

The lice were all that remained of a fantastic research laboratory where
Weigl had devised his vaccine with an almost surrealistic series of
manufacturing techniques. During World War II, Weigl’s laboratory
became the spiritual center of the city, protecting thousands of vulnerable
people who worked in it. Weigl was a bit like Oskar Schindler, the real-
life hero of Steven Spielberg’s film Schindler’s List, except that to get on
Weigl’s list, you had to strap many matchbox-size cages to your leg with a
thick rubber band. In each cage, there were hundreds of lice that fed on
your blood. The survivors of Weigl’s laboratory became famous

9
mathematicians and poets, orchestra conductors and underground
fighters.
The one who most captured my imagination was Ludwik Fleck, a
biologist in his own right, and also a philosopher of science. While
working as Weigl’s assistant, Fleck had incubated a captivating theory of
scientific knowledge and laid it out in a 1935 book, Genesis and
Development of a Scientific Fact. Today, Fleck is well known to sociologists
and historians of science. Thomas Kuhn, the famous theoretician of
knowledge who gave us the term “paradigm shift,” borrowed heavily from
Fleck’s thought in writing his 1962 classic, The Structure of Scientific
Revolutions.
Fleck’s writing drew me because of his penetrating analysis of the
human at work, his observations enlivened by clarity, humor, and
earthiness. He showered empathy upon his subjects, whether they were
medical researchers, women besotted with Parisian fashions, or medieval
astrologers. Reading Fleck gave me a sudden sense of intimacy with the
thought patterns of ancient and otherwise inscrutable people. He made
the past bubble to life by showing the integrity of its thought systems,
however wrong or bizarre they seemed to us. And he made me realize
that although we live in a world separated by an almost infinite number
of different mindsets, recognition of this fact can enable us to understand
one another. In his day job, Fleck practiced traditional scientific
reductionism, limiting the variables in order to resolve diagnostic
problems. His job was to detect the invisible particles that lurked behind
the familiar events of everyday life—the bacteria and antibodies that
helped explain a cough, a fever, a sickly child. His philosophy, by contrast,
did just the opposite: it cast a familiar light on the arcane thought
patterns of the strangers who surround us in the present and the past.
Fleck’s anthropological observations of science had allowed him to
rise above tragedy, to gain an almost spiritual perspective amid a storm-
tossed life. He had served the Habsburg Empire as a medical officer
during the Great War, endured anti-Semitic discrimination in the 1920s
and 1930s, then survived the Holocaust, the intrigues of postwar
communism, and the hush-hush of a Cold War bioterrorism lab in Israel.
Fleck’s medical specialty was immunology, or, as it was known in the
first part of the 20th century, serology—the changes in the blood that
helped doctors diagnose and treat infections. Blood had always been a
mysterious and therefore a symbolic substance—“a humor with
distinctive virtues,” as Mephistopheles says in Goethe’s Faust. That book
intrigued Fleck, who believed that the immunological paradigm of his

10
time—blood as a battleground in which cells and antibodies fought off
germs—was only the latest, culturally influenced understanding, one that
reflected the period’s nationalistic quarrels. He predicted that more
nuanced insights would open the way to calmer metaphors in the future.
He was right, as demonstrated by recent studies of the multifaceted role
of bacteria in our individual “microbiomes,” which show that we are
walking superorganisms whose life processes depend on interactions with
trillions of bacteria inside of us.
In short, more than just lice had drawn me to Lviv. So much decision
and thought and sacrifice had taken place here in this far-off corner of
Europe near the Carpathian Mountain chain. The city had been occupied
by ten different powers during eight decades. Its population had been
murdered and expelled by the hundreds of thousands in the 1940s, and the
remarkable accomplishments of these forgotten people had faded along
with their bones. Now I watched its streets fill with buses and trams and
with Ukrainian citizens, each individual possessing a feeling of belonging,
no doubt, though they inhabited a city formed by people of languages and
faiths that were absent now. It was as if Lviv’s human population was
fungible, its lice the only permanent colony.
The setting of this book is the fight against typhus during World War
II. Nazi ideology had identified typhus, which is spread by lice, as a
disease characteristic of parasitic, subhuman Jews. The Nazi medical
profession whipped itself into a terror of typhus and took outrageous
measures ostensibly to combat it. These included the walling in or closing
off of Jewish ghettos in cities like Warsaw, Kraków, and Lviv, assuring that
the disease would indeed spread, but only among Jews. Learned German
doctors convinced themselves that it was better to kill the Jews than to
allow them to contaminate others.
Weigl and Fleck thus found themselves fighting on two fronts. The
Third Reich kept them alive because it needed their expertise on typhus,
but in keeping them alive, the Nazis could not stop them from helping
others. Fleck and Weigl found the calm to practice medical science—and
to sabotage the goals of the oppressor. For their German bosses,
meanwhile, the fight against typhus became a theater of medicine gone
wrong. But there were degrees of wrongness and moral failure. Weigl’s
boss was a pragmatic army doctor, Hermann Eyer. Fleck, meanwhile, had
to work for the notorious Dr. Erwin Ding of the SS, who ruled over a
fiendish corner of the Buchenwald concentration camp, a laboratory of
ethical choices that would go on trial at Nuremberg.
In Lviv, there are no statues of Weigl or Fleck, and no monuments to

11
their work. The lice colony is nothing more than a scientific curiosity.
After World War II, a thick layer of neglect settled upon the subjects of
this book. What Fleck would have called the “thought collectives” of
typhus research, and those of Polish and Jewish Lviv in general, no longer
exist. This book attempts to clear away the dust and bring them back to
life.

12
 INTRODUCTION

Late afternoon at the Buchenwald concentration camp, on a

mountainside six miles northwest of Weimar, Germany. A cloudless day:
August 24, 1944. As always, the wind sweeps ceaselessly down the slope,
buffeting the barracks, the Gestapo bunker, the hospital, and the typhus
experimental station, blustering through the parade ground and the
ditches where road crews of matchstick inmates lift pick and shovel
accompanied by the capo’s cruel barking of orders. It is a hostile wind that
penetrates every fold in the clothing “as if it they had placed it there with
the purpose of making people feel miserable,” one prisoner will say. In the
winter it “seemed to come direct and unimpeded from the North Pole.” In
this, the eighth summer of the concentration camp’s existence, it tosses
grit into the eyes and mouth.
On the second story of Block 50, a stone-and-stucco building toward
the bottom of the hill upon which the camp stands, Ludwik Fleck runs
tests on a series of blood samples sent over from the experimental block.
He is one of a few dozen scientists from around Europe who have been
captured and brought to this building in Buchenwald to help the SS
produce typhus vaccines for the protection of German troops at the
eastern front. For two years the news from the front has been bad for the
Nazis, and the trenches are past lousy. Fleck is 48, a slight, myopic,
balding man with an expression of skeptical confidence. His slave scientist
colleagues respect his skilled hands and knowledge of the world that
swims under the microscope lens. So do the Nazi doctors who hold his life
in their hands. From his lab bench, copiously appointed with all the
equipment that the looted universities of Europe have to offer, Fleck can
see through a window and double barbed wire to the Little Camp, where
the truly doomed inmates live. Many of them are Jews like him,
stumbling along on skeletal legs amid the dirt, lice, and shit.
He hears the faint hum of the planes, a hopeful sound that is more
common now that the Luftwaffe, the German air force, has surrendered
the skies. Then the air-raid sirens sound and unexpectedly, the shriek of
explosions batters his eardrums and he falls, tossed to the wooden floor.
At last, vengeance and a direct hit! The force of the blast blows open
doors and shatters windows. Beakers and petri dishes tumble off

13
laboratory shelves, fire and mud and hot metal leap into the sky.
Electricity stops, silencing the lab’s centrifuges; with panicked shouts, the
SS duck and run and pitch themselves into their bomb shelters. The
prisoners, with nowhere to hide, jump into trenches at the edge of the
camp. D-day is two months past, Paris is on the verge of liberation, and SS
control of the concentration camps finally seems to be weakening. Forty
bombers of the Eighth U.S. Air Force have raided the military industries
adjacent to Buchenwald. Their primary target, the Gustloff-II factory,
where 3,500 camp inmates make carbines for the German army, lies in
rubble after the hourlong strike. A few incendiary bombs land on the
camp itself, and one sets fire to the Effektenkammer, the building where
the stolen possessions of the prisoners are washed and sorted, along with
the camp laundry. From there, the fire catches in the dead limbs of a large
gray tree called the Goethe oak.
This spot in central Germany was called the Ettersberg, after the
French, hêtre, for beech. In the early 19th century it was a wild and woolly
forest, 1,500 feet above sea level, part of a royal hunting ground where
German poets could commune with the inner Visigoth of their tree-
worshipping past. But that all changed in the summer of 1937, when the SS
bused in a hundred of its political enemies and ordered them to tear
down the trees and rip out the stumps. A concentration camp took shape
on the wind-swept slope—crude barracks and whipping posts for the
prisoners; villas, gardens, and a private zoo for the SS and their children.
The Nazis renamed the place Buchenwald—the beech wood.

14
Inmates walk in front of the Goethe oak in June 1944,
with the Effektenkammer in the background. (Photo
by Georges Angeli. Copyright Buchenwald
Gedenkstätte.)
Having done so, they maliciously removed all the beeches, but
preserved a single tree, a great oak six feet in diameter. Under this tree,
Johann Wolfgang von Goethe was said to have composed the Walpurgis
Night scene of his Faust. The camp commandant erected a bronze plaque
on the tree, an opportunity for the slaves to get a little culture while they
worked themselves to death. “Here Goethe rested,” it read, “during his
wanderings through the forest.”
As the war dragged on and the suffering of the prisoners passed beyond
any understanding, a legend began to circulate that the destruction of the
Goethe oak would augur the downfall of Germany. And so, as fire spread
from the laundry to the oak, some wondered whether deliverance was at
hand. There were many literate prisoners in the camp, but any affection
for the German Romantics had been replaced with an animal sensitivity
to signs and portents. The glowing embers reflected glee in the faces of
the inmate bucket brigades. The bombing raid had killed 600 inmates—
along with 200 SS men and their family members. But it was the symbolic
oak of Buchenwald that burned now, and not Paris. Ludwik Fleck
captured the mood later in an unpublished essay that was found among
his papers. “Die, die you beast, symbol of the German empire,” he wrote.
“Goethe? For us, Goethe doesn’t exist. Himmler killed him.”
The gift of Goethe, Germany’s best-loved poet, the flower of a
thinking, creative, generous world of art and science, came close to bitter
extinction in Nazi Germany. It survived, among other places, in the mind
of Ludwik Fleck, the Buchenwald laboratory slave, who had elaborated a
marvelous and prescient philosophy of science in the happier days of his
life. Science, he wrote, was a culturally conditioned, collective activity
bound by traditions that were not precisely logical and were generally
invisible to those who carried them out. Scientific disciplines, such as the
ones he belonged to, operated by the same arcane rules as a tribe in the
Amazon or a group of government clerks. The members of each thought
collective saw and believed what they had been trained to see and
believe. Pure thought and logic were illusions—perception was an activity
bound by culture and history.
The ideas and thinkers of the past were not wrong, Fleck wrote, but

15
they had built their ideas upon “shapes and meanings that we no longer
see.” This was not to say there was no progress, nor that one could not
distinguish good science from bad. The idea of an “Aryan” or a “class-
conscious” form of science, Fleck wrote in 1939—with Hitler and Stalin,
champion corrupters of science, preparing to pounce on his homeland
—“would be laughable if they weren’t so dangerous.” But Nazi medicine
was a thought collective with peculiar fixed ideas. Fleck, as a sociologist
of science, penetrated its weak points and put his insights to good use.
Arriving at Block 50 in Buchenwald in late 1943, he joined a group of
prisoners who were trying to grow typhus germs in the lungs of living,
immune-compromised rabbits. This was a task of great importance, for if
they could cultivate the germs, the cultures would be used to make a
vaccine, which would be immensely valuable to the German military. As
long as the lab was contributing to the immunological defense of its
soldiers, the Nazi regime would presumably keep the lab open and refrain
from murdering its staff. The boss, the SS Dr. Erwin Ding, would be happy,
too, because producing a vaccine would secure his position, keeping him
far away from battle duty at the eastern front. And after the war, he
hoped, the vaccine would win him a university professorship.
But there were problems with producing a sophisticated vaccine
within the thoroughly corrupted confines of a concentration camp.
Rickettsia, the intracellular bacteria that cause typhus, had bedeviled
biologists for decades. Indeed, their somewhat mysterious existence—for
no one knew just what they were—is one of the keys to understanding
how Fleck developed his questioning, skeptical view of science. These
germs were extremely difficult to grow artificially, though they thrived in
lice and sick people. The prisoners in Block 50 knew nothing about how
to prepare rickettsial cultures of the type one used to make vaccine. Yet
everyone was desperate for success—Ding to further his career, the
inmates in order to survive. Those in the thought collective of Block 50—
a biologist, a baker, a politician, and a physicist, among others—
convinced themselves that they were making a vaccine, and put the fluid
substance into vials that were sent off to Hamburg and Paris, where
German scientists, men of renown, responded with words of praise. How
was this possible? Like 1,000 monkeys with typewriters and time, a group
of desperate amateurs had learned how to prepare a devilishly
complicated vaccine in the space of a few months. Or had they? Fleck,
who arrived when the group was well advanced in its labors, was the only
one among them with the appropriate specialized knowledge. He was the
only one who knew whether the vaccine they were making was real.

16
 To most of the inmates of Buchenwald, in any case, the vaccine was
not the most interesting thing about the rabbits that were used to grow it.
On the night of the big raid, as the Goethe oak burned, the prisoners—
French scientists from the Pasteur Institute, tough Polish resistance men,
German Communists, Russian peasants, Dutch Jews, and Fleck—all joined
in singing the “Marseillaise.” Then they had a feast of rabbit stew.

17
 CHAPTER ONE

LICE/WAR/TYPHUS/MADNESS

Typhus is an unfamiliar disease to most everyone alive today, but it left

an indelible mark in past centuries, shaping the fate of empires from
Napoleon to Lenin. And no one who has lived through a typhus epidemic
will ever forget it. The disease is transmitted by tiny arthropods that live
snugly in the seams of warm clothing, which they do not leave, unless
evicted, except to suck blood or—when the body they occupy has gone
cold, or too hot with fever—to find a new human host. Lice have been
seen crawling as far as five feet in an hour. Head lice lay their nits, or eggs,
on hair; body lice, in underwear and shirts. The body louse evolved from
the head louse when people started wearing clothes and is distinguished
from its progenitor by its dislike for the temperatures of the head, our
hottest surface.
Humans and lice have a long, intimate relationship. In one of the
earliest sections of Exodus, Aaron “stretched out his hand with his rod and
smote the dust of the earth, and it became lice on man and on beast. All
the dust of the land became lice throughout all the land of Egypt.”
Expressions such as “nitwit” (for those who feel dull, sullen, and “lousy,”
after scratching their infected or allergic skin through sleepless nights)
and “nitpicking” attest to this, as does the “fine-toothed comb” we use to
examine things with care. It has even been hypothesized that the seven-
day week and the Sabbath arose in recognition of body louse reproductive
patterns—for if clothes are changed each week consistently, a person’s lice
and their eggs will die. Yet until a few centuries ago, people in colder
climates nearly always carried lice in their clothes and rarely bathed. An
account of the 12th-century funeral of Thomas à Becket notes that as his
body cooled, the vermin living in the archbishop’s many layers of clothing
began to crawl out and “boiled over like water in a simmering cauldron,
and the onlookers burst into alternate weeping and laughter.”
In the modern world, though, the body louse is the louse of refugees,
soldiers, and other desperate people. Typhus epidemics occur when a
population is at the end of its tether. Starvation, cold, fear, and
exhaustion are the normal prerequisites. Typhus corresponds with social

18
collapse. Typhus “will continue to break into the open,” wrote Hans
Zinsser, author and Harvard typhus researcher, “whenever human
stupidity and brutality give it a chance.” By the time Zinsser wrote these
lines in his famous book Rats, Lice and History, published in 1935, typhus
was a distant memory to most Americans and Western Europeans, who
were too clean for permanent louse infestation. Yet there were parts of
the world where it was still an acute danger. At the end of World War I,
the worst epidemic in history swept across Russia from Siberia all the way
through Poland, causing 30–40 million cases of disease, and killing
perhaps three million people. It was in the anteroom of this great
catastrophe that Dr. Rudolf Weigl and his assistant Ludwik Fleck earned
their stripes as typhus researchers. Working on the basis of new evidence
that lice were the vectors of the disease, Fleck and Weigl were on the
cutting edge of scientific efforts to tame it.
When the Austro-Hungarian Empire called up its male subjects to
fight in 1914, Weigl was 31 years old, Fleck just 18. Both left their homes in
the city of Lviv—which was known to the Poles as Lwów, to German
speakers as Lemberg—to become medics in the kaiser’s army. After some
training in Vienna, they quickly joined the fight against typhus, which
they encountered mostly in Russian prisoner-of-war camps in Bohemia
and in western Galicia—around the cities of Lwów, Tarnów, and Przemy
l. From 1917 to 1921, Weigl was in charge of a military laboratory—at first
under the Habsburgs, and from 1919 for the Polish state—in Przemy l,
which straddled the San River. This was a fortress town, now located at
the border between Poland and Ukraine, and by some cultural maps a
dividing line between Eastern and Western Europe. The Przemy l
complex of forts, the third largest in Europe in 1914, fell to the czar’s army
in March 1915, after a six-month siege that led to starvation among the
poor Jews who lived there. It was retaken three months later, then lost its
strategic significance and became something of a warehouse and a way
station for troop movements and a center of military medicine, including
a modern microbiological laboratory.

Rudolf Stefan Weigl was born in 1883 in P erov, a picturesque Moravian

town now located in the Czech Republic, and was the child of ethnic
Germans. His father, who designed and produced vehicles of various sorts,
died after crashing a large-wheeled bicycle of his own invention when
Weigl was seven. His mother remarried a few years later to a Polish
schoolteacher named Józef Trojnar. The family moved frequently from
town to town until Trojnar became director of a middle school in Stryj, a

19
wealthy town south of Lwów. The marriage was a happy one, and Rudolf,
his older brother, Friedrich, and sister, Lilly, grew up in an atmosphere in
which Polish language and culture predominated. After passing his
examinations, Weigl enrolled at the University of Lwów, where in 1907 he
received his doctoral degree under the zoologist Józef Nusbaum-
Hilarowicz, a leading Polish proponent and translator of Darwin’s ideas.
In the waning years of the Habsburg realm, the monarch had granted
Polish autonomy to Galicia, a district stretching from Kraków in the west
to east of Lwów and including areas of plains, forests, and mountains. The
majority of peasants in the countryside were Ukrainians, the cities
inhabited mostly by Poles and Jews. Those who chose to assimilate often
learned Polish, the language of government and culture. This contrasted
with the Russian- and Prussian-occupied areas of Poland, where
assimilated Jews tended to speak German. Perhaps because of the light
hand of the Austrian kaiser, the anti-Semitism and ethnic conflict that
would characterize Poland following its independence in 1919 were not as
close to the surface in wartime Galicia. Anti-Semitism was evident in the
professions, but had not been codified, and was not universal. Thus Weigl
was simultaneously a Czech, an Austrian, and a Pole, while his doctorate
adviser, Nusbaum-Hilarowicz, was a Jew who had decided to accept a
Catholic baptism in 1907, viewing it as a necessary step to achieve
promotion to full professor. Weigl’s boss in the military service, Filip
Pincus Eisenberg, was also Jewish. A Pasteur Institure–trained
bacteriologist, Eisenberg ran a laboratory that was as multiethnic as the
empire it served. In 1919, Weigl hired Fleck, who had begun his studies of
medicine in Lwów before the war, as his assistant in Przemy l. Fleck was
also from an assimilated background. He was the son of Sabina
Herschdörfer and of Maurycy Fleck, a craftsman with socialist tendencies
who owned a small house-painting business. Though not rich, the Fleck
parents were ambitious for their children and sent them to Polish rather
than Hebrew high schools, with hopes of offering them a way into the
mainstream of Polish society. Maurycy earned enough money to send
Fleck and his two sisters, Antonina and Henryka, to university. While
Fleck earned his doctorate at Lwów University under Weigl, the girls
studied arts and pedagogy in Vienna.
Eisenberg’s expertise was microscopy, and he was skilled at identifying
bacteria in their confusingly variable forms. Weigl, who was extremely
adept in the laboratory arts, had already invented a device for improving
microscopic lenses—a secondary focus adjustment knob. Fleck would also
gain a reputation as a razor-sharp practitioner. In visible terms, the three

20
of them represented the evolution of fashions in facial hair. Eisenberg was
balding, with a long beard of the type seen on the waistcoat-with-tails-
wearing professors in movies like The Cabinet of Dr. Caligari and The Blue
Angel. From his 20s, Weigl had sported a distinctive goatee in the manner
o f The Three Musketeers, and sartorially he favored open-necked, wide-
collared shirts. Fleck liked to dress in neatly pressed suits. He was clean-
shaven, increasingly the style as the century went on, in part because of
the nascent popular obsession with germs, which were thought to favor
beards over smooth skin (an idea encouraged by Gillette and other razor
makers).
Their prey, typhus, was an extremely difficult organism to understand
and manipulate. Most successful human pathogens are relatively benign.
Cold viruses, to give a classic example, spread far and wide because the
humans they infect remain hardy enough to distribute them among their
fellow men and women. Malaria doesn’t kill mosquitoes, and Borrelia
bacteria, the cause of Lyme disease, harm neither the deer tick nor the
deer. Over time, pathogenic organisms generally become less virulent, or
they fade away, or have limited success. (Or, as in the case of HIV and
tuberculosis, they infect slowly, giving the patient plenty of time to
transmit the germ before becoming incapacitated.) The deadly Ebola
virus sowed terror when it appeared in Africa in the 1980s, but has proven
of little global significance because it infects and kills quickly, before the
patient has time to spread it efficiently. This trait is characteristic of new
pathogens that haven’t yet adapted to their hosts. At the other end of the
spectrum is, for example, Streptococcus pyogenes, known also as Group A
strep, which can cause strep throat, toxic shock, rheumatic fever, and
scarlet fever, but usually colonizes, quietly, the throats of healthy three-
to five-year-old children—at least 15 percent of whom harbor the
organism in any given year.
Rickettsia prowazekii, according to this logic, must be a young disease,
for it has not “learned” to occupy a sturdy ecological niche. Experts
believe that American natives may have transmitted the disease to
Spanish colonists in the 16th century, although some argue it was already
present in Europe. R. prowazekii has definite shortcomings. Although it
generally kills fewer than 20 percent of the humans it infects, leaving
plenty of others alive to transmit it, typhus relies upon a single avenue—
the louse—for its spread. And lice not only spread typhus—they are its
victims. Sick insects can transmit the disease to humans for up to 10 days.
Then they die, and they do not pass along the disease to their eggs. When
there are no typhus patients around to sicken the lice, they stop

21
transmitting the disease. The end of a typhus epidemic should thus mean
the end of typhus. However, the germ has a few more survival tricks. First,
a contaminated louse’s feces contain high concentrations of R. prowazekii
and remain infectious for several months. More importantly, human
typhus survivors sometimes maintain latent infections for years. Just as a
case of chicken pox in childhood can reappear as shingles in old age,
typhus patients sometimes experience recurrences as their immune
systems weaken. An American physician, Nathan Brill, first discovered
such cases among Eastern European immigrants in New York’s Lower East
Side in 1913. They seemed especially common among elderly men and
women mourning the death of a spouse, which led Brill to call it
“bereavement disease.” Hans Zinsser isolated the organism and
indentified it as typhus. As old typhus patients die off, Brill-Zinsser
disease becomes increasingly rare. But a senior with Brill-Zinsser who
became lousy could infect his or her lice, and thus begin the cycle once
again. This mechanism keeps typhus alive between epidemics. And if, as
scientists believe, these are the only ways that typhus spreads, then the
disease will disappear from earth when the last person who ever had it
passes away.
A century ago, typhus’s unique life patterns posed a thorny challenge
to researchers. How were they to maintain a steady supply of the
organism for study? It was hard to keep typhus bacteria alive in artificial
cultures or in the bodies of mice or guinea pigs; there was no way to infect
lice with typhus other than to feed them on the bodies of people sick with
the disease. But patients were generally available only during typhus
epidemics. Even the wobbly medical ethics of those days forbade
intentionally infecting people with typhus. After many discussions with
Eisenberg about this problem, Weigl hit upon an idea in 1916. Out of
concern for his assistant’s future career, Eisenberg had been urging Weigl
to drop typhus research and concentrate on cholera, an organism that was
easier to culture and grow.
“Tell me, Sir, where are you going to get the cultures?” Eisenberg
asked. “You won’t have access to the typhus organism until you have
patients. And you won’t get patients when there is no disease. So how is
this going to work?”
Weigl thought for a moment, then with characteristic earthiness
replied, “Well, if we can’t get the louse to eat the germs, we’ll stick them
up its ass.”
Eisenberg did not understand and was not amused. Weigl told him,
“Have a look.”

22
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*** START OF THE PROJECT GUTENBERG EBOOK THE ORIGIN OF

VERTEBRATES ***
Transcriber's A few typographical errors have been corrected.
note: They appear in the text like this, and the
explanation will appear when the mouse pointer
is moved over the marked passage.

THE

ORIGIN OF VERTEBRATES

BY

WALTER HOLBROOK GASKELL

M.A., M.D. (CANTAB.), LL.D. (EDIN. AND McGILL UNIV.); F.R.S.; FELLOW OF TRINITY
HALL AND UNIVERSITY LECTURER IN PHYSIOLOGY, CAMBRIDGE; HONORARY FELLOW
OF THE ROYAL MEDICAL AND CHIRURGICAL SOCIETY; CORRESPONDING MEMBER
OF THE IMPERIAL MILITARY ACADEMY OF MEDICINE, ST. PETERSBURG, ETC.

LONGMANS, GREEN, AND CO.

39 PATERNOSTER ROW, LONDON
NEW YORK, BOMBAY, AND CALCUTTA

1908
All rights reserved
 CONTENTS

PAGE
Introduction 1

CHAPTER I
The Evidence of the Central Nervous System
Theories of the origin of vertebrates
—Importance of the central
nervous system—Evolution of
tissues—Evidence of Palæontology
—Reasons for choosing
Ammocœtes rather than
Amphioxus for the investigation of
this problem—Importance of larval
forms—Comparison of the
vertebrate and arthropod central
nervous systems—Antagonism
between cephalization and
alimentation—Life-history of
lamprey, not a degenerate animal
—Brain of Ammocœtes compared
with brain of arthropod—Summary 8

CHAPTER II
The Evidence of the Organs of Vision
Different kinds of eye—Simple and 68
compound retinas—Upright and
inverted retinas—Median eyes—
Median or pineal eyes of
Ammocœtes and their optic ganglia
—Comparison with other median
eyes—Lateral eyes of vertebrates
compared with lateral eyes of
crustaceans—Peculiarities of the
 lateral eye of the lamprey—
Meaning of the optic diverticula—
Evolution of vertebrate eyes—
Summary

CHAPTER III
The Evidence of the Skeleton
The bony and cartilaginous skeleton
considered, not the notochord—
Nature of the earliest cartilaginous
skeleton—The mesosomatic
skeleton of Ammocœtes; its
topographical arrangement, its
structure, its origin in muco-
cartilage—The prosomatic skeleton
of Ammocœtes; the trabeculæ and
parachordals, their structure, their
origin in white fibrous tissue—The
mesosomatic skeleton of Limulus
compared with that of
Ammocœtes; similarity of position,
of structure, of origin in muco-
cartilage—The prosomatic skeleton
of Limulus; the entosternite, or
plastron, compared with the
trabeculæ of Ammocœtes;
similarity of position, of structure,
of origin in fibrous tissue—
Summary 119

CHAPTER IV
The Evidence of the Respiratory Apparatus
Branchiæ considered as internal 148
branchial appendages—Innervation
of branchial segments—Cranial
region older than spinal—Three-
 root system of cranial nerves:
dorsal, lateral, ventral—Explanation
of van Wijhe's segments—Lateral
mixed root is appendage-nerve of
invertebrate—The branchial
chamber of Ammocœtes—The
branchial unit, not a pouch but an
appendage—The origin of the
branchial musculature—The
branchial circulation—The branchial
heart of the vertebrate—Not
homologous with the systemic
heart of the arthropod—Its
formation from two longitudinal
venous sinuses—Summary

CHAPTER V
The Evidence of the Thyroid Gland
The value of the appendage-unit in 185
non-branchial segments—The
double nature of the hyoid
segment—Its branchial part—Its
thyroid part—The double nature of
the opercular appendage—Its
branchial part—Its genital part—
Unique character of the thyroid
gland of Ammocœtes—Its structure
—Its openings—The nature of the
thyroid segment—The uterus of the
scorpion—Its glands—Comparison
with the thyroid gland of
Ammocœtes—Cephalic generative
glands of Limulus—Interpretation
of glandular tissue filling up the
brain-case of Ammocœtes—
Function of thyroid gland—Relation
 of thyroid gland to sexual functions
—Summary

CHAPTER VI
The Evidence of the Olfactory Apparatus
Fishes divided into Amphirhinæ and
Monorhinæ—Nasal tube of the
lamprey—Its termination at the
infundibulum—The olfactory organs
of the scorpion group—The
camerostome—Its formation as a
tube—Its derivation from a pair of
antennæ—Its termination at the
true mouth—Comparison with the
olfactory tube of Ammocœtes—
Origin of the nasal tube of
Ammocœtes from the tube of the
hypophysis—Direct comparison of
the hypophysial tube with the
olfactory tube of the scorpion
group—Summary 218

CHAPTER VII
The Prosomatic Segments of Limulus and its
Allies
Comparison of the trigeminal with the 233
prosomatic region—The prosomatic
appendages of the Gigantostraca—
Their number and nature—
Endognaths and ectognath—The
metastoma—The coxal glands—
Prosomatic region of Eurypterus
compared with that of Ammocœtes
—Prosomatic segmentation shown
by marks on carapace—Evidence of
 cœlomic cavities in Limulus—
Summary

CHAPTER VIII
The Segments belonging to the Trigeminal
Nerve-Group
The prosomatic segments of the
vertebrate—Number of segments
belonging to the trigeminal nerve-
group—History of cranial segments
—Eye-muscles and their nerves—
Comparison with the dorso-ventral
somatic muscles of the scorpion—
Explanation of the oculomotor
nerve and its group of muscles—
Explanation of the trochlear nerve
and its dorsal crossing—
Explanation of the abducens nerve
—Number of segments supplied by
the trigeminal nerves—Evidence of
their motor nuclei—Evidence of
their sensory ganglia—Summary 257

CHAPTER IX
The Prosomatic Segments of Ammocœtes
The prosomatic region in 286
Ammocœtes—The suctorial
apparatus of the adult Petromyzon
—Its origin in Ammocœtes—Its
derivation from appendages—The
segment of the lower lip or the
metastomal segment—The
tentacular segments—The tubular
muscles—Their segmental
arrangement—Their peculiar
innervation—Their correspondence
 with the system of veno-pericardial
muscles in Limulus—The old mouth
or palæostoma—The pituitary
gland—Its comparison with the
coxal gland of Limulus—Summary

CHAPTER X
The Relationship of Ammocœtes to the most
Ancient Fishes—the Ostracodermata
Cephalaspis—Ammocœtes only living
representative of these ancient
fishes—Formation of cranium—
Closure of old mouth—Rohon's
primordial cranium—Primordial
cranium of Phrynus and Galeodes
—Summary 326

CHAPTER XI
The Evidence of the Auditory Apparatus and
the Organs of the Lateral Line
Lateral line organs—Function of this 355
group of organs—Poriferous sense-
organs on the appendages in
Limulus—Branchial sense-organs—
Prosomatic sense-organs—
Flabellum—Its structure and
position—Sense-organs of
mandibles—Auditory organs of
insects and arachnids—Poriferous
chordotonal organs—Balancers of
Diptera—Resemblance to organs of
flabellum—Racquet-organs of
Galeodes—Pectens of scorpions—
Large size of nerve to all these
special sense-organs—Origin of
parachordals and auditory capsule
 —Reason why VIIth nerve passes
in and out of capsule—Evidence of
Ammocœtes—Intrusion of
glandular mass round brain into
auditory capsule—Intrusion of
generative and hepatic mass round
brain into base of flabellum—
Summary

CHAPTER XII
The Region of the Spinal Cord
Difference between cranial and spinal 385
regions—Absence of lateral root—
Meristic variation—Segmentation of
cœlom—Segmental excretory
organs—Development of nephric
organs; pronephric, mesonephric,
metanephric—Excretory organs of
Amphioxus—Solenocytes—
Excretory organs of Branchipus and
Peripatus, appendicular and
somatic—Comparison of cœlom of
Peripatus and of vertebrate—
Pronephric organs compared to
coxal glands—Origin of vertebrate
body-cavity (metacœle)—
Segmental duct—Summary of
formation of excretory organs—
Origin of somatic trunk-
musculature—Atrial cavity of
Amphioxus—Pleural folds—Ventral
growth of pleural folds and somatic
musculature—Pleural folds of
Cephalaspidæ and of Trilobita—
Meaning of the ductless glands—
Alteration in structure of excretory
 organs which have lost their duct
in vertebrates and in invertebrates
—Formation of lymphatic glands—
Segmental coxal glands of
arthropods and of vertebrates—
Origin of adrenals, pituitary body,
thymus, tonsils, thyroid, and other
ductless glands—Summary

CHAPTER XIII
The Notochord and Alimentary Canal
Relationship between notochord and
gut—Position of unsegmented tube
of notochord—Origin of notochord
from a median groove—Its function
as an accessory digestive tube—
Formation of notochordal tissue in
invertebrates from closed portions
of the digestive tube—Digestive
power of the skin of Ammocœtes—
Formation of new gut in
Ammocœtes at transformation—
Innervation of the vertebrate gut—
The three outflows of efferent
nerves belonging to the organic
system—The original close
contiguity of the respiratory
chamber to the cloaca—The
elongation of the gut—Conclusion 433

CHAPTER XIV
The Principles of Embryology
The law of recapitulation—Vindication 455
of this law by the theory advanced
in this book—The germ-layer
theory—Its present position—A
 physiological not a morphological
conception—New fundamental law
required—Composition of adult
body—Neuro-epithelial syncytium
and free-living cells—Meaning of
the blastula—Derivation of the
Metazoa from the Protozoa—
Importance of the central nervous
system for Ontogeny as well as for
Phylogeny—Derivation of free-
living cells from germ-cells—
Meaning of cœlom—Formation of
neural canal—Gastrula of
Amphioxus and of Lucifer—
Summary

CHAPTER XV
Final Remarks
Problems requiring investigation—

Giant nerve-cells and giant nerve-

fibres; their comparison in fishes
and arthropods; blood- and lymph-
corpuscles; nature of the skin;
origin of system of unstriped
muscles; origin of the sympathetic
nervous system; biological test of
relationship.

Criticisms of Balanoglossus theory—

Theory of parallel development—
Importance of the theory
advocated in this book for all
problems of Evolution 488
 Bibliography and Index of Authors 501

General Index 517

"Go on and prosper; there is

nothing so
useful in science as one of those
earthquake
hypotheses, which oblige one to face
the possibility that the solidest-
looking
structures may collapse."

Letter from Prof.

Huxley to
the Author. June
2, 1889.

THE

ORIGIN OF VERTEBRATES

INTRODUCTION

In former days it was possible for a man like Johannes Müller to be a

leader both in physiology and in comparative anatomy. Nowadays all
scientific knowledge has increased so largely that specialization is
inevitable, and every investigator is confined more and more not
only to one department of science, but as a rule to one small portion
of that department. In the case of such cognate sciences as
physiology and comparative anatomy this limiting of the scope of
view is especially deleterious, for zoology without physiology is dead,
and physiology in many of its departments without comparative
anatomy can advance but little. Then, again, the too exclusive study
of one subject always tends to force the mind into a special groove—
into a line of thought so deeply tinged with the prevalent teaching of
the subject, that any suggestions which arise contrary to such
teaching are apt to be dismissed at once as heretical and not worthy
of further thought; whereas the same suggestion arising in the mind
of one outside this particular line of thought may give rise to new
and valuable scientific discoveries.

Nothing but good can, in my opinion, result from the incursion of the
non-specialist into the realm of the specialist, provided that the
former is in earnest. Over and over again the chemist has given
valuable help to the physicist, and the physicist to the chemist, so
closely allied are the two subjects; so also is it with physiology and
anatomy, the two subjects are so interdependent that a worker in
the one may give valuable aid towards the solution of some large
problem which is the special territory of the other.

It has been a matter of surprise to many how it came about that I, a

worker in the physiological laboratory at Cambridge ever since Foster
introduced experimental physiology into English-speaking nations,
should have devoted so much time to the promulgation of a theory
of the origin of vertebrates—a subject remote from physiology, and
one of the larger questions appertaining to comparative anatomy. By
what process of thought was I led to take up the consideration of a
subject apparently so remote from all my previous work, and so
foreign to the atmosphere of a physiological laboratory?
It may perhaps be instructive to my readers to see how one
investigation leads to another, until at last, nolens volens, the worker
finds himself in front of a possible solution to a problem far removed
from his original investigation, which by the very magnitude and
importance of it forces him to devote his whole energy and time to
seeing whether his theory is good.

In the years 1880-1884 I was engaged in the investigation of the

action of the heart, and the nature of the nerves which regulate that
action. In the course of that investigation I was struck by the ease
with which it was possible to distinguish between the fibres of the
vagus and accelerator nerves on their way to the heart, owing to the
medullation of the former and the non-medullation of the latter. This
led me to an investigation of the accelerator fibres, to find out how
far they are non-medullated, and so to the discovery that the rami
communicantes connecting together the central nervous system and
the sympathetic are in reality single, not double, as had hitherto
been thought; for the grey ramus communicans is in reality a
peripheral nerve which supplies the blood-vessels of the spinal cord
and its membranes, and is of the same nature as the grey
accelerators to the heart.

This led to the conclusion that there is no give and take between
two independent nervous systems, the cerebro-spinal and the
sympathetic, as had been taught formerly, but only one nervous
system, the cerebro-spinal, which sends special medullated nerve-
fibres, characterized by their smallness, to the cells of the
sympathetic system, from which fibres pass to the periphery, usually
non-medullated. These fine medullated nerves form the system of
white rami communicantes, and have since been called by Langley
the preganglionic nerves. Further investigation showed that such
white rami are not universally distributed, but are confined to the
thoracico-lumbar region, where their distribution is easily seen in the
ventral roots, for the cells of the sympathetic system are entirely
efferent in nature, not afferent; therefore, the fibres entering into
them from the central nervous system leave the spinal cord by
ventral, not dorsal roots.

Following out this clue, I then found that in addition to this

thoracico-lumbar outflow of efferent ganglionated visceral nerves,
there are similar outflows in the cranial and sacral regions, belonging
in the former case especially to the vagus system of nerves, and in
the latter to the system of nerves which pass from the sacral region
of the cord to the ganglion-cells of the hypogastric plexus, and from
them supply the bladder, rectum, etc. To this system of nerves,
formerly called the nervi erigentes, I gave the name pelvic
splanchnics, in order to show their uniformity with the abdominal
splanchnics. These investigations led to the conclusion that the
organic system of nerves, characterized by the possession of
efferent nerve-cells situated peripherally, arises from the central
nervous system by three distinct outflows—cranial, thoracico-lumbar,
and sacral, respectively. To this system Langley has lately given the
name 'autonomic.' These three outflows are separated by two gaps
just where the plexuses for the anterior and posterior extremities
come in.

This peculiar arrangement of the white rami communicantes set me

thinking, for the gaps corresponded to an increase of somatic
musculature to form the muscles of the fore and hind limbs, so that
if, as seemed probable, the white rami communicantes arise
segmentally from the spinal cord, then a marked distinction must
exist in structure between the spinal cord in the thoracic region,
where the visceral efferent nerves are large in amount and the body
musculature scanty, and in the cervical or lumbar swellings, where
the somatic musculature abounds, and the white rami
communicantes scarcely exist.

I therefore directed my attention in the next place to the structure of

the central nervous system in the endeavour to associate the
topographical arrangement of cell-groups in this system with the
outflow of the different kinds of nerve-fibres to the peripheral
organs.

This investigation forcibly impressed upon my mind the uniformity in

the arrangement of the central nervous system as far as the centres
of origin of all the segmental nerves are concerned, both cranial and
spinal, and also the original segmental character of this part of the
nervous system.

I could not, therefore, help being struck by the force of the

comparison between the central nervous systems of Vertebrata and
Appendiculata as put forward again and again by the past
generation of comparative anatomists, and wondered why it had
been discredited. There in the infundibulum was the old œsophagus,
there in the cranial segmental nerves the infraœsophageal ganglia,
there in the cerebral hemispheres and optic and olfactory nerves the
supraœsophageal ganglia, there in the spinal cord the ventral chain
of ganglia. But if the infundibulum was the old œsophagus, what
then? The old œsophagus was continuous with and led into the
cephalic stomach. What about the infundibulum? It was continuous
with and led into the ventricles of the brain, and the whole thing
became clear. The ventricles of the brain were the old cephalic
stomach, and the canal of the spinal cord the long straight intestine
which led originally to the anus, and still in the vertebrate embryo
opens out into the anus. Not having been educated in a
morphological laboratory and taught that the one organ which is
homologous throughout the animal kingdom is the gut, and that
therefore the gut of the invertebrate ancestor must continue on as
the gut of the vertebrate, the conception that the central nervous
system has grown round and enclosed the original ancestral gut, and
that the vertebrate has formed a new gut did not seem to me so
impossible as to prevent my taking it as a working hypothesis, and
seeing to what it would lead.
This theory that the so-called central nervous system of the
vertebrate is in reality composed of two separate parts, of which the
one, the segmented part, corresponds to the central nervous system
of the highest invertebrates, while the other, the unsegmented tube,
was originally the alimentary canal of that same invertebrate, came
into my mind in the year 1887. The following year, on June 23, 1888,
I read a paper on the subject before the Anatomical Society at
Cambridge, which was published in the Journal of Anatomy and
Physiology, vol. 23, and more fully in the Journal of Physiology, vol.
10. Since that time I have been engaged in testing the theory in
every possible way, and have published the results of my
investigations in a series of papers in different journals, a list of
which I append at the end of this introductory chapter.

It is now twenty years since the theory first came into my mind, and
the work of those twenty years has convinced me more and more of
its truth, and yet during the whole time it has been ignored by the
morphological world as a whole rather than criticized. Whatever may
have been the causes for such absence of criticism, it is clear that
the serial character of its publication is a hindrance to criticism of the
theory as a whole, and I hope, therefore, that the publication of the
whole of the twenty years' work in book-form will induce those who
differ from my conclusions to come forward and show me where I
am wrong, and why my theory is untenable. Any one who has been
thinking over any one problem for so long a time becomes obsessed
with the infallibility of his own views, and is not capable of criticizing
his own work as thoroughly as others would do. I have been told
that it is impossible for one man to consider so vast a subject with
that thoroughness which is necessary, before any theory can be
accepted as the true solution of the problem. I acknowledge the
vastness of the task, and feel keenly enough my own shortcomings.
For all that, I do feel that it can only be of advantage to scientific
progress and a help to the solution of this great problem, to bring
together in one book all the facts which I have been able to collect,
which appeal to me as having an important bearing on this solution.
In this work I have been helped throughout by Miss R. Alcock. It is
not too much to say that without the assistance she has given me,
many an important link in the chain of evidence would have been
missing. With extraordinary patience she has followed, section by
section, the smallest nerves to their destination, and has largely
helped to free the transformation process in the lamprey from the
mystery which has hitherto enveloped it. She has drawn for me very
many of the illustrations scattered through the pages in this book,
and I feel that her aid has been so valuable and so continuous,
lasting as it does over the whole period of the work, that her name
ought fittingly to be associated with mine, if perchance the theory of
the Origin of Vertebrates, advocated in the pages of this book, gains
acceptance.

I am also indebted to Mr. J. Stanley Gardiner and to Dr. A. Sheridan

Lea for valuable assistance in preparing this book for the press. I
desire to express my grateful thanks to the former for valuable
criticism of the scientific evidence which I have brought forward in
this book, and to the latter for his great kindness in undertaking the
laborious task of collecting the proofs.

LIST OF PREVIOUS PUBLICATIONS BY THE AUTHOR, CONCERNING

THE ORIGIN OF VERTEBRATES.

1888. "Spinal and Cranial

Nerves." Proceedings
of the Anatomical
Society, June, 1888.
Journal of Anatomy
and Physiology, vol.
xxiii.
1889. "On the Relation
between the
Structure, Function,
Distribution, and
Origin of the Cranial
Nerves; together with
a Theory of the Origin
of the Nervous
System of
Vertebrata." Journal
of Physiology, vol. x.,
p. 153.

1889. "On the Origin of the

Central Nervous
System of
Vertebrates." Brain,
vol. xii., p. 1.

1890. "On the Origin of

Vertebrates from a
Crustacean-like
Ancestor." Quarterly
Journal of
Microscopical Science,
vol. xxxi., p. 379.

1895. "The Origin of

Vertebrates."
Proceedings of the
Cambridge
Philosophical Society,
vol. ix., p. 19.
 1896. Presidential Address to
Section I. at the
meeting of the British
Association for the
Advancement of
Science in Liverpool.
Report of the British
Association, 1896, p.
942.

1899. "On the Meaning of the

Cranial Nerves."
Presidential Address
to the Neurological
Society for the year
1899. Brain, vol. xxii.,
p. 329.

A series of papers on "The Origin of Vertebrates, deduced from the

study of Ammocœtes," in the Journal of Anatomy and Physiology, as
follows:—

1898. Part I. "The Origin of

the Brain," vol.
xxxii., p. 513.
" II. "The Origin of
the Vertebrate
Cranio-facial
Skeleton," vol.
xxxii., p. 553.
" III. "The Origin of
the Branchial
Segmentation,"
 vol. xxxiii., p.
154.
1899. " IV. "The Thyroid, or
Opercular
Segment: the
Meaning of the
Facial Nerve,"
vol. xxxiii., p.
638.
1900. " V. "The Origin of
the Pro-otic
Segmentation:
the Meaning of
the Trigeminal
and Eye-
muscle
Nerves," vol.
xxxiv., p. 465.
1900. " VI. "The Old Mouth
and the
Olfactory
Organ: the
Meaning of the
First Nerve,"
vol. xxxiv., p.
514.
1900. " VII. "The Evidence of
Prosomatic
Appendages in
Ammocœtes,
as given by the
Course and
Distribution of
the Trigeminal
Nerve," vol.
xxxiv., p. 537.
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