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Perspectives in Physiology
David H. Evans
Pioneers in Cell
Physiology:
The Story
of Warren and
Margaret Lewis
Published on behalf of The American Physiological Society

by Springer
This fascinating series seeks to place biomedical science inside a greater historical
framework, describing the main pathways of development and highlighting the
contributions of prominent investigators. This book series is published on behalf
of the American Physiological Society by Springer. Access to APS books published
with Springer is free to APS members.
David H. Evans
Pioneers in Cell Physiology:

The Story of Warren
and Margaret Lewis
David H. Evans
Department of Biology
University of Florida
Gainesville, FL, USA
ISSN 2625-2813 ISSN 2625-2821 (electronic)

ISBN 978-3-031-11893-7 ISBN 978-3-031-11894-4 (eBook)
https://doi.org/10.1007/978-3-031-11894-4
Jointly published with The American Physiological Society
© The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature Switzerland
AG 2022
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The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
Preface
During the writing of a book about the Mt. Desert Island Biological Laboratory,1 I
discovered that Warren and Margaret Lewis had a major impact on that laboratory
and the emerging field of cell physiology during the early and mid-twentieth century.
With scientific or personal connections to such luminaries as Hans Spemann,
Thomas Hunt Morgan, Ross G. Harrison, Leonor Michaelis, and Joseph Needham,
they were at the dawn of cell biology as it evolved from static anatomy to functional
physiology. Warren had formal training in William Osler’s medical school at Johns
Hopkins and was certainly the accepted leader, but Margaret (who left a Ph.D.
program to marry Warren) was the one who brought tissue culture to their research
program, for which they were best known. Her collaborations with an array of
nascent research leaders led to ground-breaking publications in hematology and
cancer causes and treatment. Throughout their long careers (which spanned over 50
years), they collaborated and also worked independently, producing a substantial
body of research literature and a library of videos of cells moving, dividing, and
engulfing their surrounding medium. This is the story of their personal and profes-
sional lives.
Gainesville, FL, USA David H. Evans
1
D H Evans, Marine Physiology Down East: the Story of the Mt. Desert Island Biological
Laboratory (New York, NY: Springer New York, 2015), pgs. 151–152.
v
Introduction
At a dinner in 1955, celebrating the lives and scientific work of Warren and Margaret
Lewis, a parade of internationally known colleagues offered their thoughts about the
couple:
We remember lunch in the laboratory; the happy conversation; the early lesson of the risk of
loose statements (and not only biological ones—has anyone ever exploited reference books,
including the Encyclopaedia Britannica, as has Dr. Warren Lewis with such devastating
effects on the inaccurate, but so good humouredly); the risk of burning the toast or getting an
unpopular cheese when it was one’s turn to do the chores. All these, and many other,
memories remain with us. But most of all the pleasure of Dr. Warren in photographing a
living egg of the pride of Dr. Margaret in thriving tissue culture. Together and alone you
have advanced biological knowledge, and are universally admired for the manner in which
you have done so. (Dixon Boyd, M.D. (and Mrs. Boyd), School of Anatomy, Cambridge,
England)
I shall never forget the morning you came in and discovered that my partner and I, who
were then dissecting ‘lowers’ and had opened the abdominal cavity, had nonchalantly
severed both vagi as they came through the diaphragm and had not bothered to work out
their distribution at all. Your withering scorn, expressed in no uncertain terms, made us both
wish that the ground beneath us would open up and swallow us whole. I was therefore all the
more surprised the following autumn when you asked me to be a student demonstrator in
gross anatomy. (Alan M. Chesney, M.D., Dean Emeritus, Johns Hopkins University Med-
ical School)
Some of us have had the great good fortune of having known and worked with you
personally and therein have benefited immeasurably from your great understanding and
extraordinary humanity. Truly, the lives and careers of those who had had such enviable
opportunity bear the indelible stamp of your influence so strikingly characterized by its
warmth, its simplicity, its friendliness and its penetrating critique. Influence such as
yours. . .is indeed immortal and we are truly glad for it. (Wiley D. Forbus, M.D., Duke
University School of Medicine)
One of my earliest memories of Woods Hole, in the twenties, is of you two walking arm
in arm, in the evening, out along the Nobska Point Road. I asked someone who you were,
and was answered in a manner which made it plain that you were something very special.
One of my earliest memories of this medical school is of you two presiding over your
laboratories in the New Hunterian, handling your cultures with loving care and with the
touch of magicians. (Allan L. Grafflin, M.D. Johns Hopkins School of Medicine)
vii
viii Introduction
I am delighted to have the opportunity of sending. . .a message of warmest greetings to

Dr. Warren H. Lewis and Mrs. Margaret Lewis upon the occasion of his 85th birthday. As
one whose work has lain in the borderline between experimental morphology and biochem-
istry, the contributions of the Lewises have been familiar to me for more than thirty years,
and one of the happiest recollections of my wife and myself concerns our visits to their home
by the edge of the water at Salsbury Cove. Representatives of the great generation of
American biologists, long may they flourish! (Joseph Needham, DPhil., Cambridge
University)
Your capacity of identifying problems which cried out for study and of devising
exquisitely suitable methods of approach to their solution proves your right to be included
in that small and select group of great leaders in the field of cell biology. Your quiet never-
ceasing thoroughness, the endless reliance upon your own hands and minds and your
apparent disdain of applause and public recognition have set examples, the influence of
which cannot be over-stated. Your world of biological science, deeply in your debt, is now,
in congratulating you, really thankfully congratulating itself for its possession of you
through all these years. (Alfred N. Richards, M.D. University of Pennsylvania)
These warm remembrances show the personal and professional esteem of the
Lewises. Their story defines the early history of biomedical research on the structure
and function of living cells, which they so carefully cultured in their unassuming
laboratories in Baltimore and Salisbury Cove.
Acknowledgments
This book could not have been written without the list of publications for Warren in
Cormer’s biography,2 as well as the list of Margaret’s publications (as well as other
documents) in the Lewis family archives in Salisbury Cove. Using these lists, and
with access to all the publications through the University of Florida’s Interlibrary
Loan system, I was able to download and read each of their publications. And the
ubiquitous Wikipedia provided access to readable explanations of biological terms,
as well as numerous biographies of scientists who were associated with Warren and
Margaret. The Lewises’ daughter Margaret Nast’s reminiscences in the Centennial
Book about the MDIBL3 provided insights into the Lewis family and their lives in
Maine. Dr. Maggy Myers (granddaughter) and Margaret Lewis (great-granddaughter
and fourth generation Margaret) kindly provided personal insights and numerous
documents, and Margaret played a significant role in what became Chaps. 1 and 2.
Without their help, this book would not have been possible.
Finally, I could not have made it through this last of my books without the love,
support, and encouragement of my wife, Jean. She has “been there” for more than 60
years.
2
G.W. Corner, “Warren Harmon Lewis, 1870-1964; a Biographical Memoir.” National Academy of
Sciences, Biographical Memoirs 1967 (1967): 321–58.
3
M N Lewis, W R Lewis, and J L Myers, “Growing Up with the Lab,” in A Laboratory by the Sea.
the Mount Desert Island Biological Laboratory, ed. F N Epstein (Rhinebeck, NY, 1998), 73–85.
ix
Contents
1 Warren Lewis: Early Life, Educational Background, and Early

Research . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
2 Margaret Reed: Early Life, Educational Background, and Early
Research . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
3 Early Collaborative Research . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
4 Move to the Carnegie Institution . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
5 Research and Family Life in the Early 1920s . . . . . . . . . . . . . . . . . 61
6 Research in the Late 1920s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
7 Warren’s Research in the 1930s . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
8 Margaret’s Research in the 1930s . . . . . . . . . . . . . . . . . . . . . . . . . . 121
9 Impact, Accolades, and a Cottage in Maine . . . . . . . . . . . . . . . . . . . 135
10 Research After Warren’s “Retirement” . . . . . . . . . . . . . . . . . . . . . 147
11 Final Years: The End of Research, But Not Honors . . . . . . . . . . . . 165
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189
xi
Chapter 1
Warren Lewis: Early Life, Educational
Background, and Early Research
Warren Harmon Lewis was born in Suffield, CT on June 17th, 1870 to John Lewis
and Adelaide Eunice Harmon, the first of their three children. The Lewis family
moved to Chicago 7 months later, where John began his law practice. Warren’s
brother John and sister Helen were both born in Chicago, in 1872 and 1876,
respectively. John was a partner in the law firm Tuley, Stiles, and Lewis, and the
family lived in Oak Park, an affluent suburb just to the west of Chicago.
Warren attended the Oak Park public schools as a child, doing well in his courses.
In 1885, when he was in the eighth grade, his scholarship average was 91, and his
teacher noted his deportment as “good.” His grades remained mainly in the 90s and
his deportment “good” and “fair,” through his time at Oak Park Public High School.
He attended the Chicago Manual Training School for his junior and senior year,
where he continued to do well in all his courses: Math, Science, and English. He
graduated in 1889. His scrapbooks1 record a popular young person, filled as they
were with calling cards and invitations to receptions and birthday parties. In his high
school years he attended many programs: musicals and orchestras. It was as a young
person that he became interested in botany. His family would recall that he was given
a copy of Asa Gray’s2 Manual of Botany as a young teen, and spent his high school
years collecting plants that he diligently pressed and kept organized in a chest.3
The family history portions of this chapter were written with the help of the Lewis’s great
granddaughter, Margaret Myers. She is the daughter of the Lewis’s grandson, John (son of
Jessica Lewis Myers, the youngest daughter of Warren and Margaret).
1
The scrapbooks are in the Lewis family archives, in their summer cottage on Spruce Point,
Salisbury Cove, ME.
2
Gray, considered “the most important American botanist of the 19th century.” See: https://en.
wikipedia.org/wiki/Asa_Gray
3
Letter from the Lewis’s daughter, Jessica Helen Lewis Myers to George Corner, dated October
11, 1965. Corner was Warren’s biographer for the National Academy of Sciences (see Footnote
14, below). In Lewis family archives.
© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 1

D. H. Evans, Pioneers in Cell Physiology: The Story of Warren and Margaret Lewis,
Perspectives in Physiology, https://doi.org/10.1007/978-3-031-11894-4_1
2 1 Warren Lewis: Early Life, Educational Background, and Early Research
Warren entered the University of Michigan in 1890. He may have been attracted
to UM because of the legacy of Asa Gray, who had been the first faculty member
appointed to the Botany department. Warren’s family seemed positive about his
decision to enroll and his ability to succeed. “Should discouragement come to you
never give up,” his maternal grandmother, Caroline (who still lived in Suffield, CT),
wrote, “persevere and all will come out right...I don’t think you really need caution
in regard to your goaheaditveness [sic]. I think it already implanted in your makeup,
an inherited gift [that] goes a good way back. . .”4 In his first semester, Warren took
French, German, Math, and systematic Zoology. He had a membership at the
University Lawn Tennis Association and continued to fill his time with concerts
and other events. His professors looked upon him fondly. A math instructor called
him “one of the very best students in my classes.” And, perhaps unsurprisingly, he
excelled in botany, as well as other areas of science.5 In October of 1893, Frederick
Charles Newcombe, an instructor in botany with a particular interest in plant
physiology, wrote that Warren was “regarded by all the instructors as one of the
best in the class, and showed marked ability, doing all his work thoroughly.”6 He
branched out from Botany, focusing also on Biology. In January of 1894, Warren
borrowed $200 from his father to purchase a microscope. The agreement specified
that he would pay back the money “on demand.”7
Warren also came under the influence of Jacob Reighard, who had returned to his
alma mater in 1886, after graduate work at Harvard and 2 years as a private tutor and
a year of high school teaching, not an uncommon pathway for a young zoologist in
the late nineteenth century. By Warren’s senior year, Reighard was a Professor of
Animal Morphology.8 After graduation in 1894, Warren remained at UM, as an
Assistant in Zoology, presumably under Reighard’s tutelage. Two years later, in
1896, Warren entered the Johns Hopkins University School of Medicine, in only the
fourth class of students. The medical school had been established in 1893, 17 years
after the university had been founded with a bequest from the estate of the American
philanthropist and abolitionist Johns Hopkins. In addition to Hopkins, the medical
school was also underwritten by several wealthy daughters of Baltimore’s business-
men, who insisted that the new medical school be open to students of both sexes,
making the new Hopkins program the first co-ed medical school in the USA.9 The
new medical school gained immediate eminence because four outstanding physi-
cians were recruited at the outset: William Henry Welch (Dean), William Osler
(medical practice), William Stewart Halsted (surgery), and Howard Atwood Kelly
4
Letter from Caroline to Warren, October 26, 1890. In Lewis family archives.
5
WHL’s scrapbook.
6
Ibid.
7
Ibid.
8
A Franklin Shull, “Jacob Ellsworth Reighard,” Science (New York, NY) 95, no. 2466 (1942):
344–46.
9
A brief history of the Johns Hopkins University School of Medicine can be found at: https://en.
wikipedia.org/wiki/Johns_Hopkins_School_of_Medicine
1 Warren Lewis: Early Life, Educational Background, and Early Research 3
(gynecology). The four are immortalized in John Singer Sargent’s The Four Doc-
tors, commissioned by Mary Elizabeth Garrett, one of the founding donors of the
medical school.10
Because of his background in morphology, Warren entered the Anatomy Depart-
ment, led by Franklin P. Mall, who had also been educated at the University of
Michigan, earning an MD in 1883.11 While doing postdoctoral research in Leipzig,
Germany, Mall started a collection of human embryos, which became the major
resource for his subsequent anatomical studies, as well as Warren Lewis’s early
research.12 Upon joining Mall’s laboratory, Warren must have immediately encoun-
tered Mall’s two junior faculty associates: Charles R. Bardeen and Ross
G. Harrison,13 both of whom would go on to make major contributions to anatomy
and embryology in the early twentieth century. Immediately after graduation in
1900, Warren joined Mall’s department as an Assistant, along with medical class-
mates Florence R. Sabin and John B. MacCallum. Each member of this group
(except MacCallum, who died at an early age), in later years, became President of
the American Association of Anatomists, and four of them (Bardeen, Harrison,
Sabin, and Lewis) were elected to the US National Academy of Sciences.14
Within a year, Warren had published his first paper, on the anatomy and devel-
opment of the human pectoralis major muscle. Using tissue from cadavers “from the
dissecting room, from bodies embalmed with the carbolic acid derivative,” he was
able to unravel the “peculiar twist in the sternocostal portion of the pectoralis major
muscle,” as well as delineate its development by dissecting some of the human
embryos that Mast had collected.15 This collection also provided the embryos for
Warren’s second publication that year. Working with Dr. Bardeen, he described the
10
http://jssgallery.org/Paintings/The_Four_Doctors.htm. To read more about the four founding
physicians, see: https://www.hopkinsmedicine.org/about/history/history5.html. Osler is probably
the best remembered, because he pioneered what became known as internal medicine, outlined in
his 1892 text, Principles of Internal Medicine and also established the first medical residency
program. For an excellent biography, see: William Osler: A Life in Medicine, by Michael Bliss.
11
As was common in the nineteenth century, Mall started his research career in Europe, studying
morphology in Heidelberg and Leipzig (where he met Welch). After a fellowship with Halstead at
Johns Hopkins, Mall held academic positions at Clark University and the University of Chicago,
before accepting the chair of the Department of Anatomy at Hopkins in 1893. See: https://embryo.
asu.edu/pages/franklin-paine-mall-1862-1917, as well as https://en.wikipedia.org/wiki/Franklin_P.
_Mall
12
For a discussion of the acquisition and importance of this collection, see: F P Mall, “The Value of
Embryological Specimens,” Maryland Med J XL, no. 3 (1898): 29–33 and F P Mall, “A Contri-
bution to the Study of the Pathology of Early Human Embryos,” Johns Hopkins Hospital Reports
IX (1900): 1–68.
13
See: https://en.wikipedia.org/wiki/Charles_Russell_Bardeen and https://en.wikipedia.org/wiki/
Ross_Granville_Harrison
14
G.W. Corner, “Warren Harmon Lewis, 1870-1964; a Biographical Memoir.” National Academy
of Sciences, Biographical Memoirs 1967 (1967): 321–58.
15
W H Lewis, “Observations on the Pectoralis Major Muscle in Man.,” Bull Johns Hopkins Hosp
12 (1901): 172–77.
development of the limbs, body-wall, and back in the early human embryo, using
serial sections embedded in wax plates, a technique which had been perfected in
Mall’s laboratory.16 They found that “during the first two months of embryonic life,
therefore, are developed the rudiments of the muscles, nerves, blood-vessels, and
skeletal structures characteristic of the back, the body-wall and the limbs. Adult
conditions are reached by an increase in size and complexity of the various organs
and by relative shifting of parts.” The paper was published as the first report in the
first volume and number of the American Journal of Anatomy, which had been
founded by Mall and three colleagues the year before “to collect into one place, and
present in a worthy manner, the many researches from our investigators, now
scattered through many publications at home and abroad.” The plates at the end of
the Bardeen and Lewis paper presented exquisite drawings of the reconstructions of
the serial plates and gross dissections of the embryos (e.g., Fig. 1.1).
This first paper with Bardeen was followed up by a second report, published in
early 1902, on the development of the human arm, again using Mall’s human
embryo collection.17 Once again, the drawings and diagrams of the dissection
were extremely detailed, but artistic (Fig. 1.2). Since Warren was the only author,
one must conclude that the drawings were his. Warren’s detailed study of human
development culminated in a comprehensive review, published in 1904 in the
multivolume Reference Handbook of the Medical Sciences, which “embrac[ed] the
entire range of scientific and practical medicine and allied science.”18
Warren spent the summers of 1895 and1901 at the Marine Biological Laboratory
at Woods Hole, MA. The MBL was the descendent of the short-lived marine station
on Penikese Island (one of the Elizabeth Islands, near Woods Hole), overseen by
Harvard’s Louis Agassiz,19 probably the leading biologist of the period. Another
MBL precursor was the Annisquam Laboratory (north of Boston), started by
Alpheus Hyatt, a student of Agassiz’s. In 1888, Hyatt moved to Woods Hole and
served as the first President of the MBL’s Corporation. Within a decade, the
Laboratory became a center for the study of the biology and physiology of marine
organisms, particularly development of invertebrates, with such luminaries as
Edmund Beecher Wilson, Thomas Hunt Morgan, Edwin Grant Conklin, Ross
Granville Harrison, and Jacques Loeb.20 During the 1901 summer at the MBL,
Warren worked with Loeb,21 studying the effect of the very poisonous compound
16
C H Bardeen and Lewis W H, “Development of the Limbs, Body-Wall and Back in Man,” Am J
Anat 1, no. 1 (1901): 1–36.
17
W H Lewis, “The Development of the Arm in Man.,” Am J Anat 1, no. 2 (1902): 145–183.
18
W H Lewis, “Development of Foetus,” in Reference Handbook of the Medical Sciences, ed. A H
Buck, 2nd ed., vol. 8, (New York, 1904), 450–57.
19
See: https://en.wikipedia.org/wiki/Louis_Agassiz
20
J Maienschein, “Agassiz, Hyatt, Whitman, and the Birth of the Marine Biological Laboratory,”
Biol Bull 168, no. 3 (June 1985): 26–34.
21
Loeb was an established physiologist at this point, with an appointment at the University of
Chicago. He was best known for his research on animal tropisms, toxic and antitoxic effects of ions,
Fig. 1.1 Drawing of a lateral view of human embryo, age about 5 weeks. Screen shot of Plate
V from: C H Bardeen and Lewis W H, “Development of the Limbs, Body-Wall and Back in Man,”
Am J Anat 1, no. 1 (1901): 1–36
potassium cyanide on the life-span of unfertilized sea urchin eggs.22 This was
Warren’s first foray into cell and developmental biology, a significant change from
mammalian morphology. Loeb had previously found that shed urchin eggs died
quickly, while fertilized eggs started to divide and develop into embryos. He
concluded from these studies that there are “two kinds of processes going on in
the egg—a mortal process and a second which leads to cell division and further
development. The latter process inhibits or modifies the mortal process.” The
research with Warren set out to determine if the mortal process could be delayed,
by “agencies which inhibit catalytic phenomena without permanently altering the
artificial parthenogenesis, and heteromorphosis (“the replacement of an injured or removed organ

by a different organ”). See: https://en.wikipedia.org/wiki/Jacques_Loeb
22
J Loeb and W H Lewis, “On the Prolongation of the Life of the Unfertilized Eggs of Sea-Urchins
by Potassium Cyanide,” Am J Physiol 6 (1902): 305–17.
Fig. 1.2 Drawing of medial view of arms of two human embryos, age 5–7 weeks. Screen shot of
Plate II from: Lewis W H, “Development of the Arm in Man,” Am J Anat 1, no. 2 (1902): 145–183
constitution of living matter.” Since potassium cyanide was known to inhibit “a

number of enzymatic processes,” they exposed unfertilized urchin eggs to various
concentrations of the poison before fertilizing them. They found that control eggs
(no poison) could reach the pluteus stage of development if fertilized within 23 h;
eggs developed for longer periods would not develop to the pluteus stage and “as a
rule, the eggs form a sticky and discolored mass.” Eggs kept in cyanide solutions
(0.001% was optimal), however, could develop to the pluteus stage even after 112 h
of exposure. Higher concentrations of cyanide precluded any development. Lack of
oxygen did “not prolong, or prolongs but little, the life of fertilized eggs.” They also
considered the nature of the “mortal process” of the normal egg. What was going on
that resulted in death if no fertilization took place? They concluded that “no definite
answer is possible at present,” but posited that it might be “self-digestion” or, since
the “nucleoproteids” of the cell nuclei may be “oxidizing agents,” as the cell divides
and the contents of the original nucleus “are scattered throughout the cell. It is easily
conceivable that this periodic spreading or mixing of the contents of the nucleus and
the cells may modify the chemical processes in the egg and check the mortal
processes.” This hypothesis is especially interesting because the importance of
nucleoproteids (now termed nucleoproteins), such as proteins associated with
DNA, was not known in the early twentieth century. We now know, of course,
that nuclear DNA controls not only cellular division, but also the biochemical
metabolism of cells.23
The summer working with Loeb at the MBL, must have shifted at least some of
Warren’s interests to more experimental approaches, because many of his subse-
quent publications examined processes associated with changing morphology
(development), rather than morphology itself.
Returning to vertebrate development and morphology, and working in the labo-
ratory of Professor Moritz Nussbaum in Bonn, Switzerland (presumably sometime
in 1902), Warren showed definitively that the “wandering pigment cells” in the optic
cup of the chick, as well as the pupillary muscle, are of ectodermal origin and,
therefore, “must modify our idea of a rigid specificity of the outer germ layer.”24
This was one of the early suggestions that adult tissues, such as the vertebrate eye,
might be composed of cells from more than one germ layer (ectoderm, mesoderm,
endoderm), a concept that was becoming a major research topic in developmental
biology, and of Warren’s research.
By the spring of 1903,25 Warren was studying the interactions between the
developing amphibian eye cup and the overlying epidermis (skin). Warren’s
research had been prompted by an earlier (1901) study by Hans Spemann,26 which
had shown that the developing eye of the frog, Rana fusca, did not produce a lens if
the early optic cup was destroyed by electrocautery. In addition, if the optic cup had
not been destroyed completely, the lens formed only if the cup rudiment touched the
23
For a brief introduction to the structure and function of DNA, see: https://biologydictionary.net/
dna/ or https://en.wikipedia.org/wiki/DNA
24
Warren Harmon Lewis, “Wandering Pigmented Cells Arising From the Epithelium of the Optic
Cup, with Observations on the Origin of the M. Sphincter Pupillae in the Chick,” Am J Anat 2, no.
3 (July 1, 1903): 405–16.
25
Warren was promoted to Assistant Professor of Anatomy at Johns Hopkins Medical School that
year. A year later, he was promoted to Associate Professor, a rate of academic advancement much
faster than normal.
26
See: https://en.wikipedia.org/wiki/Hans_Spemann
epidermis. These experiments suggested that lens development was dependent on

optic cup development.27 In a later review of this work, Viktor Hamburger trans-
lated,28 from the original paper, Spemann’s “intent” for this ground-breaking study:
The complex apparatus of the vertebrate eye originates by a sequence of developmental
processes which. . .are interlocked spatially and temporally. The following is an experimen-
tal contribution to the question of whether these developmental processes proceed depen-
dently or independently of each other; that is, whether their spatial and temporal
coordination is guaranteed by a causal interaction or by a harmony which dates back to
earlier stages, and perhaps to the egg.
Warren Lewis had read Spemann paper (while in Nussbaum’s lab in 1902?) and
decided to approach the question a bit more directly the next year. He wrote: “By the
use of the binocular microscope one can make minute dissections of the living
amphibian embryo and can remove various organs, transplant them or alter the
normal relations, and so alter the influences they exert on each other. We may thus
determine certain correlations necessary to normal development.”29 This was one of
the first demonstrations of the importance of microdissection under a binocular
microscope. Warren was able to actually remove the optic cup (vesicle) before the
lens had started to differentiate from the ectoderm (skin) in the frog Rana sylvatica.
Like Spemann, he found that if the optic vesicle failed to regenerate, no lens
developed. If the vesicle “regenerates sufficiently to come in contact with the skin
a lens will form.” If the regenerated eye cup “is deeply buried. . .a lens does not
form.” The next two series of experiments were ground-breaking. Warren found that
if the optic cup was transplanted to a more caudal portion of the embryo, a lens
developed if the transplanted optic cup touched the skin, but did not if the optic cup
was buried more deeply and did not touch the skin. And he could show that skin
transplanted from the abdomen of the frog to over the eye cup could develop a lens
only if the transplanted skin touched the developing eye cup. He concluded that “the
lens is absolutely dependent for its origin on the influence of the optic vesicle on the
ectoderm” and “there is no predetermined area of ectoderm which must be stimu-
lated in order that a lens may arise.” The abstract ends with one of the earliest
descriptions of what came to be known as “embryonic induction”:
We have here one tissue influencing another during the course of development, and from this
a new structure, the lens, arises. It seems likely that there is a definite chemical reaction
between certain substances of the optic vesicle and certain substances of the ectoderm cells,
which results in the formation of new substances within the lens cells, and that these
substances give to the lens its peculiar characters and mode of development.
27
H Spemann, “Uber Correlationen in Der Entwicklung Des Auges,” Verhandl Anat Ges 15 (1901):
61–79.
28
V Hamburger, The Heritage of Experimental Embryology: Hans Spemann and the Organizer,
(New York, Oxford: Oxford University Press, 1988). 196 pgs.
29
W H Lewis, “Experimental Studies on the Development of the Eye in Amphibia,” Am J Anat
Suppl 3 (1904): xiii–xv. For a short discussion of embryonic induction, see https://encyclopedia2.
thefreedictionary.com/embryonic+induction
Fig. 1.3 Drawing of transverse section of an amphibian embryo in which the early optic cup had
been surgically removed on the left side and then reinserted away from the ectoderm. Two days
later, the embryo was sacrificed and the development of the lens on the left compared with that in
the normal eye on the right side. It is clear that neither a regenerated eye cup (rg) nor the
transplanted eye cup (tr) had stimulated the production of a lens, as seen on the right side. Screen
shot of Fig. 3 in W H Lewis, “Experimental Studies on the Development of the Eye in
Amphibia. I. on the Origin of the Lens. Rana palustris.,” Am J Anat 3 (1904): 505–36
In a subsequent publication (also published in 1904), Warren gave a much more

complete description, with illustrations, of this work.30 He noted that he had used
two species of frogs (R. sylvatica and R. palustris) to do the transplantation studies,
to take advantage of the different pigmentation in their skin. In addition to describing
the extirpation (Fig. 1.3) and transplantation (Fig. 1.4) experiments outlined in the
1904 abstract, he demonstrated that “the all-important influence of the optic vesicle
30
W H Lewis, “Experimental Studies on the Development of the Eye in Amphibia. I. on the Origin
of the Lens. Rana palustris.,” Am J Anat 3 (1904): 505–36.
Fig. 1.4 (a) Drawing of a transverse section of an amphibian embryo in which the early optic cup
had been surgically removed on the left side and then transplanted into the flank of the embryo,
caudal from the cephalic region. Two days later the eye cup had developed further and induced the
formation of a small lens between it and the ectodermal tissue, as long as the original cup tissue was
in contact with the ectodermal tissue. Screen shot of Fig. 11 in W H Lewis, “Experimental Studies
on the Development of the Eye in Amphibia. I. on the Origin of the Lens. Rana palustris.,” Am J
Anat 3 (1904): 505–36. (b) Drawing from a follow-up experiment, where the transplanted optic cup
was not in contact with the ectoderm. In this case, no lens is induced. Screen shot of Fig. 26 in W H
Lewis, “Experimental Studies on the Development of the Eye in Amphibia. I. on the Origin of the
Lens. Rana palustris.,” Am J Anat 3 (1904): 505–36
is brought out most forcibly by the demonstration of the power it possesses to cause
the formation of a lens from ectoderm taken from over the abdomen of a different
species.” He concluded that “it would seem evident that chemical changes have
taken place within these lens-forming ectodermal cells, and that the optic vesicle
cells by some means have been able to change the chemical nature of these cells, new
substances being formed which are now peculiar to the groups of cells that show
such a modified development. It would seem probable that the modified develop-
ment was directly due to these new chemical substances.” In other words, the
developing optic cup cells were somehow able to control the differentiation of
non-specific skin cells to produce the lens.
The importance of Spemann’s and Lewis’s studies cannot be overstated. They
were the first to show that the development of specific vertebrate tissues and organs
is not predetermined in the egg but is the result of cell and tissue interactions during
the early stages of development. This ideas of “induction” and “organizers” were the
bases for extensive studies in embryology during the next half-century.31 In Ham-
burger’s words: “Spemann and Lewis deserve credit for the first experimental
documentation of a case of embryonic induction, which turned out later to be an
important and widespread mechanism in epigenetic development.” Subsequent
research has shown that, at least in some species, lens development is not controlled
by the optic cup, so this specific case of induction is not universal, but the general
phenomenon of tissue interactions during embryonic development is. For this work,
as well as many subsequent studies, Hans Spemann was awarded the Nobel Prize in
1935, the first embryologist to be so honored.32 Surprisingly, the chemical nature of
the “organizers” is still disputed (and may depend on the specific case of induction),
but are most likely protein products of DNA.33
Next, Warren applied his new technique of microsurgery under a binocular
microscope to a study of the development of the amphibian cornea, the clear area
of skin that overlies the eye.34 Spemann had shown that the optical clearing of the
developing skin to form the cornea was dependent on the presence of both the optic
cup and the lens (see footnote 28). Warren, however, determined that, at least in the
salamander Amblystoma, “corneal clearing will occur over a naked lens or over the
optic cup without the lens, provided the lens or cup are close to the overlying
ectoderm.”35 Moreover, he was able to show that “the cornea will form from
ectoderm other than that which normally gives rise to the cornea. . .[and] that even
after the cornea is formed it will disappear almost completely if the optic cup and
lens is entirely removed, without injury to the overlying cornea.” He concluded this
work with the following thoughts about the underlying causes of corneal differen-
tiation and tissue interactions during development in general:
It seems very probable that the optic vesicle brings about lens formation through a specific
influence. The cornea, however, in so far as its early stages are concerned, namely, the
thinning and clearing of the skin and loss of pigment can hardly be ascribed to a specific
influence. That the mechanical pressure of the eye, or cup or lens may in some way be
accountable for the corneal changes is a possibility. I am more inclined to the view, however,
that the changes are due to another and quite different reason. The contact of either the eye or
cup or lens with the epidermal cells must of necessity alter the environment of the overlying
cells as regards their relation to the mesenchyme.36 It is possible that this exclusion from
31
Reviewed by: Hamburger, The Heritage of Experimental Embryology: Hans Spemann and the
Organizer; (New York: Oxford University Press, 1988); J M W Slack, From Egg to Embryo, 2nd
ed., (Cambridge: Cambridge University Press, 1991).
32
See:
https://embryo.asu.edu/pages/hans-spemann-1869-1941, as well as: https://www.nobelprize.org/
prizes/medicine/1935/spemann/lecture/
33
Domenico Ribatti, “The Chemical Nature of the Factor Responsible for Embryonic Induction: an
Historical Overview.,” Organogenesis 10, no. 1 (January 1, 2014): 38–43.
34
See: https://en.wikipedia.org/wiki/Cornea
35
W H Lewis, “Experimental Studies on the Development of the Eye in Amphibia. II. On the
Cornea,” J Exp Zool 2 (1905): 431–46.
36
Mesenchyme is a type of connective tissue found in vertebrate embryos. See: https://en.
wikipedia.org/wiki/Mesenchyme
contact with the mesenchyme cells may be responsible for changes in the metabolism of the
epidermal cells and cause them thereby to alter their mode of development, such alterations
leading to corneal changes. That such apparently slight alterations in environment are
responsible for other important changes in the history of embryonic ectodermal cells I
think quite probable. From experiments already completed it seems highly probable that
the central nervous system is in part at least dependent for its origin and differentiation on the
difference of environment of cells which at one time possessed the possibilities of producing
either ordinary epidermal cells or of producing nerve cells.
Warren extended his studies of eye development in a paper, published in 1907,

that described further experiments dealing with the development (“induction” was
still not used) of the lens in amphibians. These studies were largely confirmatory, but
they also found that lens formation required the “continued influence of the optic
vesicle and optic-cup [and] probably only the retinal portion of the optic vesicle is
capable of stimulating lens-formation from the ectoderm.” Moreover, the size of the
lens appeared to be correlated with the length of time that the optic vesicle or optic
cup “remains in contact by its retinal layer with the growing lens-structure.” He also
proposed that the stimulus from the underlying optic cup “may be only
mechanical.”37
In December of 1905, members of the Anatomical Laboratory from Johns
Hopkins attended the annual meeting of the Association of American Anatomists
in Ann Arbor, MI and presented research results from the previous year. Warren
gave two talks, one of which described experiments on the regeneration and differ-
entiation of the central nervous system in amphibian embryos. After a fertilized egg
divides sufficiently to form a ball of cells (called a blastula), a specific portion of the
ball invaginates to form an interior cylinder of cells within the ball. The embryo is
now called a blastula, and the opening into the interior is called the blastopore.38
Warren described how cells that had been transplanted from the dorsal lip of the
blastopore into mesenchyme of a much older embryo of the frog, Rana palustris,
could differentiate “into spinal cord, notochord and muscle, indicating that the cells
of the rim of the half-closed blastopore are already predetermined.” In addition,
small pieces of early brain tissue could be transplanted into older mesenchyme and
form a “considerable portion of the brain corresponding to the region which would
normally have developed about the piece that had been cut out.” And the brain
region surrounding the area lost during transplantation would “regenerate the lost
part and a perfectly normal brain will result.”39 In a review of the discovery of what
37
W H Lewis, “Experimental Studies on the Development of the Eye in Amphibia. III. on the
Origin and Differentiation of the Lens,” Am J Anat 6 (1907): 473–509.
38
See https://en.wikipedia.org/wiki/Gastrulation for a general description of gastrulation. See also
57 s to 1:39 min in the video: https://vimeo.com/315487551
39
W H Lewis “Experiments on the Regeneration and Differentiation of the Central Nervous System
in Amphibian Embryos,” Am J Anat Suppl 5 (1906): xi. A more complete description of these
studies was published a year later: W H Lewis “Transplantation of the lips of the blastopore in Rana
palustris,” Am J Anat. 1907;7:137–143.
has been termed “Spemann’s Organizer,” Fässler and Sander40 proposed that this
study by Warren was in fact the first to describe the “organizer” that was credited to
the work of Spemann and his student Hilde Mangold.41 They wrote that, in 1927, Sir
Gavin R. DeBeer stated: “It must not be forgotten that while the detailed analysis of
these phenomena is the work of Spemann and his school, the credit of making the
first organizer graft belongs to Lewis (1907).” They noted that Warren had “grafted
the organizer from a gastrula of Rana palustris into the otic region of another embryo
in the tailbud stage, and obtained an induction of nerve tube, notochord and
somites.” They added that “In fact, during his long and distinguished career Lewis
never claimed to have discovered the organizer effect—not even when the tempta-
tion to claim this might have been greatest, namely when soon after the Second
World War he hosted a postdoctoral student from Otto Mangold’s laboratory.”42
Warren’s second presentation at that 1905 meeting (as well as one given at the
same meeting by Ross Harrison) described part of a collaboration between Warren
and Harrison on the development of the peripheral nervous system. Harrison had
become interested in the controversy about the mechanisms of formation of amphib-
ian peripheral nerves (called axons43), which go from the central nervous system out
to peripheral tissues, such as the skin, eye, stomach, and muscles. At that time, it was
unclear if axons developed from an outgrowth from the central nervous system or by
the fusion of numerous, individual cells in the periphery in a sort of chain. Harrison’s
experiments showed that axons develop from the CNS outward, not by individual
cells forming a chain.44 Warren’s careful dissection and transplantation studies of
amphibian embryos provided a system to corroborate Harrison’s findings. Warren
described experiments that demonstrated that if the nasal pit of developing
Amblystoma is transplanted to adjacent ectoderm before its nerves have developed,
it “will send out its nerve fibers between it and the ectoderm at which place there is
no mesenchyme.” If the anterior end of the brain is removed before the nerves
develop from the transplant, the nerves from the transplant will grow into the
mesenchyme that has filled the space left from the extirpated brain tissue, suggesting
that the nerve “paths. . .are in no sense predetermined.” Moreover, transplanted
pieces of the developing neural tube will send out nerves on their own, even into
40
P E Fässler and K Sander, “Hilde Mangold (1898-1924) and Spemann's Organizer: Achievement
and Tragedy,” in Landmarks in Developmental Biology -1883-1924, ed. K Sander, vol. 22, (Land-
marks in Dev Biol, 1997), 66–75. The author thanks Prof. Malcolm Maden from the University of
Florida Department of Biology for bringing the Fässler and Sander paper to his attention.
41
H Spemann and H Mangold, “Uber Die Induktion Von Embryonalanlagen Durch Implantation
Artfremder Organisatoren,” Arch Mikrosk Anat Entwicklungsmech 100 (1924): 599–638.
42
Hilde’s husband, Otto, had been a student of Spemann’s and her collaborator before her tragic
death in 1924. P E Fässler and K Sander, “Hilde Mangold (1898-1924) and Spemann's Organizer:
Achievement and Tragedy,” in Landmarks in Developmental Biology -1883-1924, ed. K Sander,
vol. 22, (Landmarks in Dev Biol, 1997), 66–75.
43
For a general description of an axon, see: https://en.wikipedia.org/wiki/Axon
44
R G Harrison, “Further Experiments on the Development of Peripheral Nerves,” Am J Anat
6 (1906): 121–31.
strange mesenchyme or the wall of the nearby pharynx. Lastly, if the brain tissue is
damaged just after the closure of the neural fold, new nerves may exit “into the
mesenchyme in various directions.” Warren suggested that “the only adequate
explanation of such phenomena seems to me to lie in the acceptance of the outgrowth
theory of the axis cylinder.”45 The next year (1907) Warren published a more
extensive paper on the outgrowth of the axis cylinder, concluding that “these
experiments furnish important evidence in favor of the out-growth theory of the
nerve fiber, and also that the fiber is a process of a nerve cell and not from a chain of
cells.”46
Because his earlier studies had shown that lens development was dependent upon
interaction with the underlying optic vesicle, while the optic vesicle itself (as well as
the notochord, muscle, and central nervous system) was “self-differentiating,”
Warren was interested in determining the developmental interactions between
other embryonic organs and their surrounding tissues. In a study also published in
1907, his microsurgical techniques demonstrated that the otic vesicle (which
becomes the ear of the adult amphibian) is indeed self-differentiating, not dependent
on interactions with nearby cells or tissues.47
That same year (1907), Warren published his final paper48 on amphibian eye
development, focusing on the “differentiation and regeneration of the eye at an
earlier stage, at about the time of or shortly after the closure and beginning fusion
of the neural folds.”49 At this stage in development, the optic vesicle “projects from
the side of the brain and produces a bulging of the ectoderm on the surface of the
embryo [and] has remarkable powers of self-differentiation.” The cavity of the optic
vesicle communicates with the brain cavity, the optic stalk has not formed, and there
is “no indication of any histological differentiation between those cells which are
destined to form the eye or the optic stalk and brain.” However, optic vesicles
transplanted to other regions of the developing embryo were able to subsequently
grow and differentiate optic tissues such as the retinal and pigment layers. These
were the same experiments that demonstrated that such transplanted optic vesicles
could “induce” the formation of ectodermal lenses and corneas. This, unlike the
developing lens and cornea, which development is dependent upon “the continued
influence of the optic vesicle for [their] origin, early growth, and differentiation,” the
optic vesicle itself is self-differentiating.
45
W H Lewis, “Experimental Evidence in Support of the Outgrowth Theory of the Axis Cylinder,”
Am J Anat Suppl 5 (1906): x–xi.
46
W H Lewis, “Experimental Evidence in Support of the Theory of the Outgrowth of the Axis
Cylinder,” Am J Anat 6 (1907): 461–471.
47
W H Lewis, “On the Origin and Differentiation of the Otic Vesicle in Amphibian Embryos,” The
Anatomical Record 1 (1907): 141–45.
48
W H Lewis, “Experiments on the Origin and Differentiation of the Optic Vesicle in Amphibia,”
Am J Anat 7 (1907): 259–77.
49
After gastrulation, the dorsal surface of vertebrate embryos forms a neural fold, which closes to
form the brain and spinal cord. See: https://en.wikipedia.org/wiki/Neural_fold, as well as minutes 1:
40 to 2:10 of the video: https://vimeo.com/315487551
During this same period, Warren’s stature in the world of anatomical research was
attested to by the publication of his extensive review of what was “the first complete
text-book of human anatomy of any considerable importance written and produced
in this country entirely by American authors,”50 George A. Piersol’s Human Anat-
omy, Including Structure and Development and Practical Considerations. The tome
was 2088 pages long and published in 1907 by Lippincott Company of Philadelphia
and London. Warren admitted that “in reviewing a book of this size. . .the reviewer
can scarcely be expected to read every page, must less attempt to indicate either all
its virtues or defects.” He noted, however, that “during the past winter I have had
frequent occasion to consult the text, which has proved excellent and concise, while
its use by a number of my students to their entire satisfaction indicates that it will
soon be widely and extensively used in this country.” Further, he suggested that the
cost of the volume ($7.50), “is less than one-half the cost of Quain’s Anatomy.51
This very remarkable price will contribute much to the success of the book.”
Warren’s major problem with the new text was with the illustrations, which “espe-
cially at the periphery, are not carefully worked out and often are mere smudges.”
And “I have often found it necessary to turn to figures in other books not because the
views are not well chosen, but because the artist has often failed to draw with the eye
of an anatomist.” Given the exacting and elegant illustrations in his own publications
(See Figs. 1.1 and 1.2), Warren was certainly a knowledgeable critic. He also
regretted that “more references to the literature have not been added. Such references
are extremely valuable to the student, and as an introductory method into the
literature serve as a distinct educational feature.” Despite these quibbles, Warren
ended the review: “The anatomy as a whole is a decided success, and the editor and
his collaborators are to be congratulated for giving us a truly first-class text-book.”
Warren’s interest in morphology and development continued in 1909–1910, with
the publication of a massive chapter on “The development of the Muscular System”
in Keibel and Mall’s Manual of Human Embryology, which must have been the
standard text on human embryology of the time.52 He also showed that the limb
muscles of the developing amphibian do not derive from the lateral myotomes but
50
W H Lewis, Review of Human Anatomy, Including Structures and Development and Practical
Considerations, edited by George A. Piersol, Philadelphia, Lippincott, 1907. Anat Rec 2 (1908):
259-287. Warren noted that the book was “a distinct step... that must be gratifying to all American
anatomists. We are no longer dependent upon England or Germany for a text-book of the first rank.”
It is important to add that Warren’s stature in the field was also shown by the fact that by the
publication of Vol 2 (1) (May, 1908) of the Anatomical Record, Warren’s name was on the
masthead, as a member of the Editorial Board.
51
Elements of Descriptive and Practical Anatomy, written by Jones Quain, was first published in
1828 in England. Subsequent editions, some edited by his brother Richard, were available until the
early twentieth century.
52
W H Lewis, “The Development of the Muscular System,” in Manual of Human Embryology,
ed. F Keibel and F P Mall, (Philadelphia, 1910), 454–522.
develop in situ.53 And he found that transplanted pieces of the embryonic amphibian
neural plate would develop normally and showed “great regenerative power.”54
Warren spent at least some of his summer research time at the Marine Biological
Laboratory at Woods Hole during this period (1904–1909, except for 1906), and he
presented one of the famous Friday Evening Lectures in 1905.55 Three publications
arose from these MBL summers. The first was a study of the experimental produc-
tion of cyclopia in the killifish, Fundulus heteroclitus.56 An earlier investigator had
found that cyclopia could be produced in this small, inshore fish by incubating the
developing embryos in a MgCl2 or Mg(NO3)2 solution in seawater.57 Warren
proposed that the cyclopian condition “can [not] be looked upon as germinal in
origin but [is] truly due to environmental conditions.” Once again using microsur-
gery, Warren found that he could reproduce Stockard’s cyclopian killifish by
insertion of a “very fine needle” through the egg membrane into the anterior end
of the embryonic shield, which forms as gastrulation takes place. He suggested,
therefore, that the chemical induction had “in some manner prevented the growth of
certain cells at the anterior end of the embryonic shield during the embryonic shield
stage.” The second study was on the early cleavage stages of the killifish. By
removing small portions of the early blastodisc, before or after cleavage, he was
able to show that there was “no definite fixed localization of organ forming sub-
stances.” In addition, he also found that “the position of the first cleavage plane is
predetermined.”58 In a subsequent study, Warren showed, in later stages of killifish
development (embryonic shield formed) “removal of any portion of the axis of the
embryonic shield results in very definite defects in corresponding parts of the
embryo. . .[and] the embryonic shield does not possess the power of regeneration
of the parts removed from the axis.” He suggested that “this method of experimen-
tation affords an opportunity for experimental work on the central nervous system by
the elimination of various portions at various stages. The development of the
vascular system can also be studied experimentally.”59
It was during the summer of 1905 at the MBL that Warren (Fig. 1.5) met Margaret
Reed, a graduate student and research associate of Thomas Hunt Morgan from
Columbia University. Their personal and scientific lives would never be the same.
53
W H Lewis, “The Relation of the Myotomes to the Ventrolateral Musculature and to the Anterior
Limbs in Amblystoma,” Anat Record 4 (1910): 183–90.
54
W H Lewis, “Localization and Regeneration in the Neural Plate of Amphibian Embryos,” Anat
Record 4 (1910): 191–98.
55
Appreciation is expressed to Ms. Jennifer Walton of the MBL Library for this information.
56
W H Lewis, “The Experimental Production of Cyclopia in the Fish Embryo (Fundulus
heteroclitus),” The Anatomical Record 3 (1909): 175–81.
57
C R Stockard, “The Question of Cyclopia, One-Eyed Monsters,” Science, New Series 28, no.
718 (1908): 455–56.
58
W H Lewis, “Experiments on Localization in the Eggs of a Teleost Fish (Fundulus heteroclitus),”
Anat Record 6, no. 1 (1912): 1–6.
59
W H Lewis, “Experiments on Localization and Regeneration in the Embryonic Shield and Germ
Ring of a Teleost Fish (Fundulus heteroclitus),” Anat Record 6, no. 1 (1912): 325–33.
Fig. 1.5 Copy of a

photograph of Warren
Lewis, taken in 1904 in
Baltimore. (Original in
Lewis family archives)
Chapter 2
Margaret Reed: Early Life, Educational
Background, and Early Research
Margaret Adaline Reed was born on November 9, 1881 to Martha (often called
Mattie) Adaline Walker Reed and Joseph Cable Reed. She was born, like her father
and older sister Jessica, in Kittanning, PA, a small town on the east bank of the
Allegheny River, 44 miles northeast of Pittsburgh. In her autobiography,1 Margaret
recorded her “uneventful” childhood in Kittanning, but did recall the Johnstown
Flood of the Conemaugh River in May of 1889, when she was 8 years old: “I went
from place to place in a row boat with my father, helping to rescue stranded people.”
The Lewis family had moved to Meyersdale, PA (near Johnstown) earlier. Years
later, Margaret’s eldest daughter (Margaret Nast, introduced later) wrote that her
mother had a happy childhood, despite having a bout with Scarlet Fever2 at a young
age, leaving her with hearing loss which worsened as she got older. Margaret Nast
also wrote3 that her mother proclaimed her intention to be a doctor at age 13 which
was not met “with wholehearted rejoicing by her parents.” Apparently, however, this
should not have come as a complete surprise, as Margaret had shown an interest in
medicine even as a young teenager, joining a Doctor Rowe “in his horse-drawn
buggy when he made country house calls.”4
Margaret did not spend much of her teenage years in Meyersdale. In 1894, Mattie
and Joseph decided that their two daughters should attend a finishing school and
chose to send the girls to the Hannah More Academy, an Episcopal boarding school
in Maryland. Margaret and Jessica, however, were apparently unhappy with that
The family history portions of this chapter was written with the help of the Lewis’s great
granddaughter, Margaret Myers. She is the daughter of the Lewis’s grandson, John (son of
Jessica Lewis Myers, the youngest daughter of Warren and Margaret).
1
Margaret Reed Lewis Autobiography, in Lewis family archives.
2
See: https://en.wikipedia.org/wiki/Scarlet_fever
3
Memoirs of Margaret Nast Lewis, in Lewis family archives.
4
Ibid.

20 2 Margaret Reed: Early Life, Educational Background, and Early Research
plan, and family legend5 has it that the night the girls were dropped off, they snuck
off campus and found their parents at their hotel. There, the girls expressed their
unhappiness with the school, and stated that they instead wished to attend the
Women’s College of Baltimore (now Goucher College6). At the time, Margaret
was 14, and Jessica 15. Joseph and Mattie did take the girls to the Women’s College,
where a counselor suggested they spend a few years at the Girls’ Latin School of
Baltimore,7 which at that time was a preparatory institution associated with and on
the campus of the Women’s College. After spending the requisite time at the
preparatory school, Margaret entered the Women’s College of Baltimore. “Then
the joy of college,” she wrote.8 “My class was 1901; we loved the old granite
buildings and church.” She graduated with a degree in math and was a member of
the Alpha Chapter of the Tau Kappa Pi Fraternity. She is remembered as “a little
lady, always daintily dressed, quiet, but a person with an unbelievable capacity for
hard work.”9 In her yearbook from the college,10 each senior was asked to imagine a
building for a “twentieth century city.” Among the schools, artistic institutions,
homes, and charitable institutes, we find Margaret’s proposed building: a “Marine
Biological Laboratory,” which she described as “one of the most notable buildings
on the waterfront, and one which attracts many students of science to the city. It is
finely equipped for scientific work, and much original investigation leading to
tactical results is pursued there. The work is characterized by enthusiasm, thorough-
ness and untiring perseverance.”11
Margaret’s choice of a “Marine Biological Laboratory” as a twentieth century
building was presumably based upon her summer of 1900 study at the Marine
Biological Laboratory at Woods Hole, MA, the start of her career in biological
research.12 She probably had been recruited to MBL by Thomas Hunt Morgan,
because she entered graduate school, and Morgan’s laboratory, at Bryn Mawr in
1901, after her second summer at the MBL.13 Morgan was soon to be a very famous
5
Ibid.
6
See: https://en.wikipedia.org/wiki/Goucher_College
7
See: https://en.wikipedia.org/wiki/Girls%27_Latin_School_of_Baltimore
8
Margaret Reed Lewis Autobiography, Op. Cit.
9
Goucher Alumnae Quarterly, 29, (1951), pgs. 16–17.
10
Goucher College Donnybrook Fair 1902 Year Book, pg. 39. In Goucher College Archives. See:
https://mdsoar.org/handle/11603/1713
11
Ibid.
12
These notes on Margaret’s schooling and early career are summarized from: M Ogilvie and J
Harvey, The Biographical Dictionary of Women in Science, vol. 2, pgs. 785-786 (New York,
London: Routledge, 2000); the Bryn Mawr Alumnae Quarterly, 1907–1909, pg. 66; M J Hogue,
Margaret Reed Lewis. Goucher Alumnae Quarterly 7 (3) (1929):49-52; as well as G W Corner,
“Warren Harmon Lewis, 1870–1964; a Biographical Memoir,” National Academy of Sciences,
Biographical Memoirs 1967 (1967), pgs. 331–332.
13
Bryn Mawr College Program of Graduate Courses, 1902, pg. 14 and 1903, pg. 15. Appreciation is
expressed to Ms. Marianne Hansen, Curator of Rare Books and Manuscripts, Bryn Mawr College
2 Margaret Reed: Early Life, Educational Background, and Early Research 21
developmental biologist and geneticist.14 Margaret remained at Bryn Mawr for

2 years, working again with Morgan at the MBL in the summer of 1902.15 She
returned to Bryn Mawr for the 1902–1903 school year, on a scholarship.16 That year,
she studied with Professor Joseph Weatherhead Warren, taking a graduate physiol-
ogy class where the topics were “Mechanics of Digestion,” and “The Knee-Jerk.”
About Margaret, Dr. Warren wrote that she “faithfully attended these courses and
manifested an industrious attention.”17
In 1903, Margaret moved to New York to begin a 3-year Lectureship at the
New York Medical College for Women; she held a similar position at Barnard
College in 1905–1907.18 It is apparent that she was still working in Morgan’s
laboratory19 during these years (he had moved to Columbia University in 1904),
and in 1907 (still without a graduate degree) she accepted the position of Instructor
in Biology and Physiology at Columbia University and remained there until 1909.
Margaret apparently returned briefly to Bryn Mawr in 1908, because she is listed as a
Fellow in Biology at Bryn Mawr during the 1908–1909 academic year.20 Early in
her time in New York, Margaret’s hometown newspaper (The Meyersdale Republic)
reported that “the young lady for the past few years has been making this line of
study her specialty and this is her reward. It is certainly a great honor to the young
lady, and her many friends in this city extend their hearty congratulations to the
young lady on her well deserved [Sic] success.”21 In her summers, Margaret traveled
extensively. She spent the summer of 1904 in the Rocky Mountains in Colorado. The
Republic reported that there she “even [entered] a gold mine and [dug] some of the
precious metal.” On her way home from Colorado, Margaret stopped in St. Louis,
for finding this resource. Margaret’s peripatetic early career is also outlined in a list of academic
positions in the Lewis family archives.
14
During his early years (1890–1900), Morgan’s research centered on invertebrate embryology and
regeneration (using tadpoles, fish, and earthworms). He was awarded the Nobel Prize in 1933 for his
work delineating the role of chromosomes in heredity, using the fruit fly Drosophila
melanogaster. See: https://en.wikipedia.org/wiki/Thomas_Hunt_Morgan
15
Morgan reported after the 1901–1902 school year that “Miss Margaret Reed has taken a course of
grad. Lectures in Regeneration with me, and has taken part of the Journal club work. She had
worked in the lab. on the Regeneration of the limbs of the crayfish and on the devel. of a whole limb
in the newt where only one of the bones of the fore-arm is present.” “Report of Professor,” in Lewis
family archives.
16
The Lantern, Bryn Mawr College, 1902 (Philadelphia: Avil Printing Company). See: https://
archive.org/details/lantern1115stud/lantern1115stud_djvu.txt.?view¼theater
17
Note in Lewis family archives.
18
Bryn Mawr College Register of Alumnae and former students, 1920, pg. 199.
19
In the Columbia University Catalog for 1906–1907, Morgan is listed as a “Professor of Exper-
imental Zoology” and Margaret is listed as a “Tutor in Zoology in Barnard College.” Barnard had
been founded as a private women’s liberal arts college in 1889, but was soon incorporated into
Columbia in as one of its four undergraduate colleges. See: https://en.wikipedia.org/wiki/Barnard_
College
20
Note in graduate record for Margaret in Bryn Mawr Archives.
21
“Secures Good Position,” The Republic, June 11, 1903.
where the Louisiana Purchase Exposition was taking place.22 There, she might have
seen some of the most exciting technological and medical advances of the day,
including the wireless telephone, automobiles and airplanes, X-ray machine, and
infant incubators, which were all on display at the exposition.23
In the summer of 1906, Margaret traveled to Europe to spend the first of three
summers there, sightseeing and doing some scientific research at the University of
Zurich. During that summer, Margaret and friends explored the Alps, duly noted by
the Republic: “When they reached the Alps, instead of going around by rail, she with
Miss Dederer and Prof. Lewis,24 accompanied by guides made the trip across on
foot.”25 This trek in the Alps was recorded by a photo, now in the Lewis family
archives (Fig. 2.1). The paper noted that this trip was “a most hardy undertaking for
women.” She wrote fondly of the trip, but seemed to find the trek difficult: “So you
see, dear,” she wrote to her parents on August 7, 1906, “it is a rather footsore little
maiden that writes, for we carried our knapsacks also, as well as climbed ten hours. It
was by far the most beautiful day I have ever known.”26
During this period, Margaret published three papers while working in Morgan’s
laboratory, two from her research at Bryn Mawr (1903 and 1904) and one from work
at Columbia (1905).27 In the first paper, from research done “at the suggestion of
Professor Morgan. . .in the winters of 1901/02 and 1902/03,” Margaret wrote about
regeneration of a whole foot in a salamander from the cut end of a leg containing
only the tibia.28 She had found that, if the surgical removal of the fibula is carefully
done, “the new part that regenerates will produce a whole new foot with five toes,
and also the distal end of the fibula. These structures are formed from material
proliferated by the cut-end of the tibia alone.” In the second paper “under the
direction of Professor Morgan. . .during the winters of 1902 and 1903,” Margaret
presented data from her study of the regeneration of the first leg of the crayfish29
asking “how the cells to form the new muscles of the regenerated leg arise, whether
22
“Miss Margaret Reed, Who Has,” The Republic, September 15, 1904.
23
See: https://en.wikipedia.org/wiki/Louisiana_Purchase_Exposition
24
As indicated in the previous chapter, it is probable that Margaret had met Warren Lewis the
previous summer (1905) at the Marine Biological Laboratory at Woods Hole.
25
“Miss Reed in the Swiss Alps,” The Republic, August 30, 1906. The paper had published two
letters that Margaret had sent to her parents.
26
“Miss Reed in the Swiss Alps,” The Republic, August 30, 1906.
27
It is interesting to note that Margaret was only 22, and a year out of college, when the first paper
was published. Morgan was not a co-author on any of the papers, which is also unusual, especially
for early papers written by a graduate student.
28
M A Reed, “The Regeneration of a Whole Foot From the Cut End of a Leg Containing Only the
Tibia,” Arch F. Entwicklingsmeschn D. Organ 17 (1903): 150–54.
29
Like many invertebrates, and some vertebrates (such as a salamander), crayfish can regenerate a
limb that has been lost. For a recent review of this interesting phenomenon, see: Sunetra Das,
“Morphological, Molecular, and Hormonal Basis of Limb Regeneration Across Pancrustacea.,”
Integr. Comp. Biol. 55, no. 5 (November 2015): 869–77, doi:10.1093/icb/icv101.
Fig. 2.1 Screen shot of a photograph of Margaret Reed and Warren Lewis in Switzerland in the
summer of 1906. (Original in Lewis family archives)
from the old muscles, which are uninjured, or from some other source.”30 She found
that “the muscles of the new leg formed from cells proliferated by the ectoderm. . .
[and] the cells to form the new nerve (or at least the nerve sheath) are also
differentiated from cells proliferated by the ectoderm.” She also used the hermit
crab in similar studies and found that two new legs could be produced if the nerve is
divided or cut in two. In the third paper, “following Professor Morgan’s suggestion,”
30
M A Reed, “The Regeneration of the First Leg of the Crayfish,” Archiv Fur
Entwicklungsmechanik Der Organismen 18, no. 3 (January 1, 1904): 307–16.
Margaret wrote about her study of early cleavage in the frog’s egg, to try to
determine if the layer of interior cells was produced by the same cleavage planes
as the surface layer, or by delamination from the outer layer of cells. She found that
“sections through the egg as late as the end of the 32-cell stage show that all the cells
are divided by cleavage-planes, which appear on the surface.” Thus, her observa-
tions “show that there is no delamination division cutting off a number of cells
parallel to the surface but that the first formed cells of the interior are produced by the
same division planes as are the cells of the surface and by their subsequent division
from the cells of the interior”31
In the summer of 1907, Margaret returned to Europe, this time taking a steamship
from New York to Naples, Italy.32 Her letters to her parents detailed her travels in
Italy, including stops in Pasestuna, Poscidonia, Pompei, Cara, Amalfi, Sorrento, and
Capri.33 The next summer, Margaret went to Berlin to study in the laboratory of Max
Hartmann and worked with his young colleague, Rhoda Erdmann.34 The Republic
reported that Margaret had written to her parents that the institution was “equipped
with everything imagineable [sic] to work with, as well as being scrupulously
clean.”35 In her own writing about her graduate work abroad, Margaret wrote that
she “profitted [sic] greatly by my association with Dr. Rhoda Erdman and Professor
Max Hartman. My how Max could sing! Dr. Erdman believed that living cells would
eventually be grown in glass.”36 She had been growing (culturing) amoebae on
agar37 substrate kept moist with a distilled water. Using Erdmann’s techniques (with
the substitution of Ringer’s/Locke’s solution38 for distilled water), Margaret cul-
tured guinea pig bone marrow cells and “found that [these cells] formed a
membrane-like growth with mitotic figures on the surface of nutrient agar.”39 This
appears to be the first successful culture of mammalian cells.40 The techniques of
producing growth of isolated cells and tissues originated in the early work
31
M A Reed, “The Formation of the Interior Cells of the Segmentation of the Frog's Egg,” Biol Bull
8 (1905): 189–89.
32
“Home News,” The Republic, July 18, 1907.
33
“Touring Land of Sunshine,” The Republic, September 12, 1907.
34
See: https://en.wikipedia.org/wiki/Max_Hartmann and https://en.wikipedia.org/wiki/Rhoda_
Erdmann
35
“Miss Margaret A. Reed Back from Germany,” The Republic, November 26, 1908.
36
37
See: https://en.wikipedia.org/wiki/Agar
38
Ringer’s Solution is a saline solution (NaCl in distilled water) with the addition of other salts,
such as K, Ca, and HCO3. It was first formulated by Sydney Ringer, a nineteenth-century British
physiologist and pharmacologist. See: https://en.wikipedia.org/wiki/Sydney_Ringer. Locke’s solu-
tion is Ringer’s solution with glucose and slightly more NaCl added.
39
There is no published record of this initial work by Margaret, but it is described briefly in the first
paragraph of a subsequent publication: M R Lewis and W H Lewis. “The Growth of Embryonic
Chick Tissues in Artificial Media, Agar and Bouillon. Bull Johns Hopkins Hosp 22 (1910):126-127.
40
M A Harvey, “Johns Hopkins--the Birthplace of Tissue Culture: the Story of Ross G. Harrison,
Warren H. Lewis and George O. Gey,” The Johns Hopkins Medical Journal 136 (1975): 142–49.
(1906–1908) of Johns Hopkins’ Ross Harrison on the mechanisms of amphibian

nerve regeneration and growth. His techniques not only provided the first evidence
that nerves grew outward from the spinal column (See previous chapter), but they
also enabled the culture of living cells outside of the amphibian body.41 By 1913, the
technique that came to be called “tissue culture” had been modified and extended to
chick and mammalian tissues, largely due to the work at The Rockefeller Institute
(now University) by Drs. Alexis Carrel42 and Montrose Burrows, who published a
series of papers in 1910–1913.43
Margaret stayed on in Europe in the fall of 1908, reporting that her work was
going “very well as far as the Pathological side is concerned,” and that she had been
asked by Professor Metchnikoff44 to visit the Pasteur Institute in Paris.45 The
Republic reported in November of 1908, in a tone of hometown pride, that “in her
studies Miss Reed has indeed made a remarkable record, and is rapidly becoming
one of the country’s leading authorities in the line of research which she is pursuing.”
The paper also reported that she was researching tuberculosis, and that she would go
back to Bryn Mawr, “where she will pursue the same work that she was engaged in
during her stay in Germany.”46 On September 9, 1908, she wrote to her parents from
Paris, presumably while visiting the Pasteur Institute, with her plans to take a
steamer back from Germany to Bryn Mawr. “I expect to be there until June,” she
wrote of the college, “but if things do not go well I will come back to Germany. The
climate drives me away now but I did like it there more than any place I have been
and I am very homesick to go back.”47
41
R G Harrison, “Further Experiments on the Development of Peripheral Nerves,” Am J Anat
6 (1906): 121–31; R G Harrison, “Observations on the Living Developing Nerve Fiber,” The
Anatomical Record 1 (1907): 116–18; R G Harrison, “Embryonic transplantation and the develop-
ment of the nervous system,” The Harvey Lectures. Series 4 (1908): 199–222.
42
In 1909, Carrel had sent his young assistant, Burrows, to Harrison (then at Yale) “to learn the
latter’s methods and adapt then to tissues of warm-blooded animals” G W Corner, “Warren Harmon
Lewis, 1870-1964; a Biographical Memoir,” National Academy of Sciences, Biographical Memoirs
1967 (1967): 321–58. Carrel received the Nobel Prize in Physiology or Medicine in 1912 “for
pioneering work in vascular suturing techniques.” He also invented the first perfusion pump (with
Charles Lindbergh) “opening the way to organ transplantation.” See: https://en.wikipedia.org/wiki/
Alexis_Carrel
43
A Carrel and M T Burrows, “Cultivation of Adult Tissues and Organs Outside of the Body,” J Am
Med Assoc LV, no. 16 (1910): 1379–80; A Carrel and M T Burrows, “Cultivation of Tissues in
Vitro and Its Technique,” J Exp Med 13, no. 3 (1913): 387–405; M T Burrows, “The Cultivation of
Tissues of the Chick-Embryo Outside the Body,” J Am Med Assoc LV, no. 24 (1910): 2057–58; M
T Burrows, “The Growth of Tissues of the Chick Embryo Outside the Animal Body, with Special
Reference to the Nervous System,” J Exp Zool 10 (1911): 63–85; M T Burrows, “The Tissue
Culture as a Physiological Method,” Trans Cong Am Physicians and Surgeons 9 (1913): 77–90.
44
Elie Metchnikoff was awarded the Nobel Prize Physiology and Medicine that year. See: https://
en.wikipedia.org/wiki/Élie_Metchnikoff
45
Letter from Margaret to Warren, Oct. 26, 1908. In Lewis family archives.
46
“Miss Margaret A. Reed Back from Germany,” The Republic, November 26, 1908.
47
“Letter from Gay Paree,” The Republic, Dec. 3, 1908.
In December of 1908, Margaret applied for the Ph.D. program at Bryn Mawr in
physiology and morphology; despite her continued work since graduating from
college, she had not yet taken a graduate degree from any other institution. In her
application, she detailed her extensive undergraduate and graduate study of biology
and zoology, as well as her proficiency in French and German. She proposed
studying Physiology primarily, with a “subordinate subject” in morphology. She
outlined the many courses she had taken with Dr. Morgan in experimental embry-
ology, cellular biology, and experimental zoology. She proposed a thesis with the
topic: “Study of the leucocytes of the Blood.” “Work for this,” she wrote on her
application, “was begun 1906 and carried on systematically ever since.”48 There are
no records of Margaret working toward a graduate degree after the 1908–1909 year.
In her brief autobiography, she wrote that “she started some research work at Bryn
Mawr College, but left it to marry Dr. Warren H. Lewis.”
Sometime between 1905 and 1908, Margaret had fallen in love with Warren
Lewis. The actual timing is difficult to confirm, but Warren and Margaret had both
been at the MBL during the summer of 1905, and they certainly were together in the
Alps in the summer of 1906 (see above). The relationship had progressed, as
Margaret wrote:
It was a snowy night in Baltimore when my friend of five years, Dr. Warren H. Lewis,
Professor of Anatomy at the Johns Hopkins Medical School and I came out of the Anatom-
ical Building. He turned, kissed me and said, “you know I love you, want to marry you and
have you bear our children.” I could not speak. Me! Me! Become a wife and mother? “Give
me time to learn.” “Yes until May” and in those happy days I learned the sweetness of love.49
And on January 10, 1909, Warren wrote to Margaret (at Radnor Hall at Bryn
Mawr, where she had returned to graduate school for 1 year):50
Margaret dear it was good to get your letter this evening and to come close in touch with you
as I read it over. These are ever wonderful days my beloved with you close in my thought
and feeling world. It is all too good and beautiful to be true, so like the things of one’s dream
world where the imagination carries one inside the deepest and truest relations with the
dream companion. And now the ideal companionship of which I have often dreamed of
seems really to be developing. But I am afraid sometimes that you will be disappointed in not
finding all the depths and understanding that you imagine lie within this personality of mine.
There is a big deep something there how much I do not know yet—it expresses itself in many
ways on the surface and is [?]51 not only through those expressions that one learns of it. I do
not isolate it from the many phases of one’s everyday living. This inside self that you value
above all else how can one know it in others or oneself ever except by outward expression,
Often unconscious though it be. I do love you dear for this inside self yet what I know for it is
learned only through your letters, our talks, and by many little observations of your ways and
work and manners. Many of these observations are perhaps unconscious, an intuitive mode
which often I think tells more than all the others. Yes it is a new world of our own that we are
48
Application in Lewis family archives.
49
50
The handwritten letter is in the Lewis family archives. At this point in time, Margaret was 27 and
Warren was 39. The letter is decidedly formal and Victorian, but very personal.
51
[?] has been inserted when the writing is unclear.
building a world without limits—if built up in true love which is more than friendship or
comradeship or emotions yet-these all form parts of it. We cannot really picture how good or
beautiful this world will be but this I feel is what happiness, real happiness, cannot come or
last without—this harmony of what you call the inside self.
Such a happy day has this one been Margaret dear with all the beautiful memoires of the
wonderful sweet maid who came to play with me for three days. I am so happy dear in my
love for you that I just want to sing and skip for joy. It was indeed a very happy time and I am
so glad that it is you dear heart who is to live this beautiful life close with me and it is so good
that you are happy with such an old fellow as I. I will try dear and learn many beautiful things
for you have much to teach me.
I wish I could make you feel dear how happy I am with you and how beautiful your
whole personality is to me.
I have felt all day the harmony and am full in flowing me with an elation of happiness and
your letter tonight my beloved breath of your own happiness from this beauty and harmony
and [?] beyond measure my own.
Last night was a beautiful dream of you I thought as I awoke from it, I would surely
remember in the morning and how bad that it has all slipped away.
The pretty “wind bells” tinkle sweetly to me as I write as they hang on the chandelier in
this [?] They fill the room with sweet thoughts of you sweet-maid.
May all the days be beautiful and happy ones for you dear and bring we two ever into a
more beautiful harmony.
And now we begin the days of work for us that are to be successful and full of happiness
to each one with deep thoughts of each other to help.
Ever, Warren
The last sentence of the letter suggests that they were about to begin their research
collaboration—which lasted until Warren’s death 54 years later. By October of
1909, they must have announced their engagement, because Warren’s father, John,
wrote to his future daughter-in-law:
Dear Margaret,
I thought I should have written you long before this but somehow there does not seem to
be much occasion for the written word. I am only less interested in Warren than you are and I
am prepared to welcome his wife most heartily into our family circle. I have longed to see
Warren happily married and I shall feel very grateful and tender towards the woman who
enables him to realize that desire. I do not mean that it should be the fault of anything I can
help if you do not like your future father in law. I most earnestly hope that you will never
regret this step you have taken and I believe you will not; I look forward to a more intimate
acquaintance with you at no very significant delay. And now and always I wish you every
joy and good tiding and most of all a happy marriage with my first-born.
Most sincerely yours,
John Lewis52
Margaret and Warren were married on May 23rd, 1910.
52
Handwritten letter in Lewis family archives.
Chapter 3
Early Collaborative Research
By the fall of 1910, Warren and Margaret had begun collaborative research, which
continued for the next 54 years. Margaret’s early experiments with culturing mam-
malian tissues in Rhoda Erdmann’s laboratory in 1908, and Warren’s close relation-
ship with Ross Harrison’s early work on culturing embryonic amphibian neurons
must have prompted them to begin a more in-depth study of the best experimental
conditions for the successful culturing of various vertebrate (and invertebrate)
tissues. It soon became clear that the actual composition of the culture medium
was of paramount importance. Erdmann and Margaret had used distilled water and
then Locke’s solution, and Harrison had initially used amphibian lymph.1 The
Lewises’ early studies, using embryonic pig bone marrow in plasma from older
pigs, produced inconclusive results. With the appearance of the Burrows and Carrel
research,2 which used clotted plasma, they “began similar studies on embryo chicks
of various ages and were able to obtain. . .very wonderful growths from the various
organs of the embryos.” They initially used Harrison’s hanging drop method3 but
found that they could produce much more extensive cultures in Petri dishes, “the
diameter of these growths sometimes exceeding 1 centimeter.” The Petri dish
method offered “certain advantages for experimentation on the growth, which can
1
See: M R Lewis and W H Lewis, “The Growth of Embryonic Chick Tissues in Artificial Media,
Agar and Bouillon,” Bull Johns Hopkins Hosp. 22. (1911): 126-127; R G Harrison, “Observations
on the Living Developing Nerve Fiber,” The Anatomical Record 1 (1907): 116–18.
2
A Carrel and M T Burrows, “Cultivation of Adult Tissues and Organs Outside of the Body,” J Am
Med Assoc LV, no. 16 (1910): 1379–80.
3
From Harrison (1907; Op.Cit.): “After carefully dissecting it out, the piece of tissue is removed by
a fine pipette to a cover slip upon which is a drop of lymph freshly drawn from one of the lymph-
sacs of an adult frog. The lymph clots very quickly, holding the tissue in a fixed position. The cover
slip is then inverted over a hollow slide and the rim sealed with paraffin. When reasonable aseptic
precautions are taken, tissues will live under these conditions for a week and in some cases
specimens have been kept alive for nearly 4 weeks. Such specimens may be readily observed
from day to day under high magnifying powers.” See also: https://microbeonline.com/procedure-
hanging-drop-method-test-bacterial-motility/

30 3 Early Collaborative Research
be subjected to the influence of various fluids for varying lengths of time.” The fact
that they produced similar growths from a variety of tissues (e.g., liver, intestine,
heart, brain, kidney, adrenal, muscle, stomach, thyroid, retina, and cornea from chick
embryos from 10 to 21 days old) suggested that “the growing tissues are only such as
are common to the various organs, namely, connective tissues of various kinds and
endothelium of blood vessels.” The found that the clotted plasma could be replaced
by a mixed salt solution4 and that cultured intestinal tissue continued “regular
peristalsis for over 48 hours,” and other cultured cells showed “active growth and
migration of cells.” The cells were “active, sending out and retracting their pseudo-
podia,5 and sometimes moving slowly apparently on the underside of the
coverslip.”6
It is important to note that Margaret was the first author of this first joint paper
with Warren, indicating her importance in the research, not to mention her presumed
major role in the actual writing of the manuscript. The next paper produced during
this first year of collaboration also had Margaret as first author. It was a logical
continuation of these studies: “having found that tissues from chick embryos grow
readily in artificial media, nutrient agar and bouillon, the next step was the attempt to
cultivate such tissues in media all the constituents of which are known. . .It is hoped
that an artificial medium will be found as satisfactory as the plasma, for the
advantages are obvious if one can work with a known medium in the investigation
of the many new problems, which suggest themselves.”7 Using laborious, ionic-
substitution protocols, they found that, while tissues (liver or liver plus intestine,
kidney, heart, or spleen) could not grow in NaCl-free media, they grew “readily” in
media that did not contain either CaCl2, KCl, or NaHCO3 (Fig. 3.1). They concluded
that “NaCl is apparently the most essential salt and although the others are not
necessary the maximum growth was attained when all four were used.” The Lewises
were apparently aware of the importance of the osmotic pressure of the
media vs. that of the cells, but made no effort to control this important variable,
writing: “It is quite remarkable that cells will grow in such widely different solutions
where osmotic pressure must vary considerably. It will be interesting to determine
the osmotic pressures of these solutions.”
The Lewises summarized this stage of what they described as “a new epoch in
experimental anatomy”:
The introduction by Harrison of the important method of the study of growth of small pieces
of living tissues of cold-blooded animals transplanted into lymph and the later extension of
4
Initially, they formulated a medium containing NaCl, KCl, CaCl2, NaHCO3, H2O, agar, and
bouillon, but later studies used a medium without the agar.
5
Pseudopodia are temporary extensions of the cell membrane (with contain cytoplasm) of cells,
used primarily for movement or feeding. See: https://en.wikipedia.org/wiki/Pseudopodia
6
This and the preceding quotes are from: Lewis and Lewis (1911, Bull Johns Hopkins Hosp.
22, Op. Cit.).
7
M R Lewis and W H Lewis, “The Cultivation of Tissues From Chick Embryos in Solutions of
NaCl, CaCl2, KCl and NaHCO3,” Anat Record 5 (1911): 277–93.
3 Early Collaborative Research 31
Fig. 3.1 Photomicrograph

(X 60 diameters) of cells
growing out from a 48-h
culture of chick liver tissue.
Screen shot of Fig. 11 in M
R Lewis and W H Lewis,
“The Cultivation of Tissues
From Chick Embryos in
Solutions of NaCl, CaCl2,
KCl and NaHCO3,” Anat
Record 5 (1911): 277–93
this method by Burrows and Carrel to warm-blooded animals by transplanting into plasma
marks the beginning of a new epoch in experimental anatomy.
The result of the observations given [in our work], that embryonic chick tissue will grow
in various salt solutions, further broadens the field of work in that tissues can be studied in
media all the constituents of which are known and the reaction of the cells to different
substances can be more definitely determined.8
Thus, during the first year of their marriage, Margaret and Warren had begun
ground-breaking collaboration, showing that cells could be grown outside of an
organism’s body using only a mixed salt solution. As we shall see, the Lewises
continued to play a major role in the perfection and evolution of cell and tissue
culture—techniques that remain important in biomedical research over 100 years
8
M R Lewis and W H Lewis, “The Cultivation of Tissues in Salt Solutions,” Jama 56, no.
24 (1911): 1795–96.
later.9 And, they had started a family, with the birth of their first daughter, Margaret
Nast Lewis, on August 21, 1911.10
Margaret and Warren continued their studies of chick tissues in culture during
1912. Following up on the ground-breaking studies of the culture of amphibian
nerves (in lymph or plasma) by Harrison and Burrows (see footnotes 1 and 2), they
reported on the culture and growth of chick sympathetic nerves in the defined salt
solutions they had previously described.11 In their introductory statement, they
described the “advantages of a salt solution medium over that of lymph or plasma”:
One great advantage which the cultivation in salt solution prosses over the method of
Harrison and Burrows depends on the fact that the outgrowing nerve fibers creep along
the underside of the coverslip and seem to stick closely to it. . .thus the fibers are for the most
part all in one plane [of] focus and can be followed continuously even with the oil immersion
without difficulty. Another advantage is that in the fixing and staining the fluid salt solutions
are much simpler to deal with than the more complicated plasma or lymph clot with its fibrin
threads which often take the stain and obscure the picture presented by the growing tissues.
The most important advantage however consists in the fact that we are dealing with solutions
of known chemical constitution and that our picture is uncomplicated by structures except
those which have grown out from the original piece.
In this study, they demonstrated that the cultured nerve fibers developed rapidly,
attaining a length of over 1 mm, and showed “the characteristic amoeboid nerve ends
and are rich in lateral branches and true anastomoses with neighboring
fibers. . .appear to be granular in structure [and] react to the iron haematoxylin12
very much as do the chromosomes.” They extended this study to other tissues of the
embryonic chick in a subsequent paper, because their earlier studies had shown that
“such growths continued for several days but were never as extensive either in
9
Margaret’s and Warren’s importance in the early research on tissue culture is described in three
excellent reviews of this field: M A Harvey, “Johns Hopkins--the Birthplace of Tissue Culture: the
Story of Ross G. Harrison, Warren H. Lewis and George O. Gey,” The Johns Hopkins Medical
Journal 136 (1975): 142–49; H Landecker, The Lewis Films: Tissue Culture and “Living Anat-
omy,” 1919-1940, In: J Maienschein, M Glitz and G E Allen, “Centennial History of the Carnegie
Institution of Washington, Vol. V The Department of Embryology,” Cambridge University Press
(2004): 117-144; H Landecker, Culturing Life. How Cells Become Technology, Cambridge, MA:
Harvard University Press (2007), 276 pgs. Harvey’s review presents the importance of tissue
culture: “This phenomenal research tool is of major importance in many fields of scientific
investigation including cytology, histology, embryology, virology, cancer research, radiobiology,
molecular biology and biochemical genetics. The development of important new knowledge
through the use of tissue culture techniques has become so extensive that it has been
designated by many as the greatest technical advance in medical science since the invention
of the compound microscope.” Emphasis added by this author.
10
Margaret Nast would become a scientist in her own right, graduating from Goucher College in
1931 with an AB in physics, followed by a Ph.D. in physics from Johns Hopkins in 1937. She was a
member of the Harvard College Observatory from 1961 to 1986 and died at the age of 101 in
2012. See: https://news.harvard.edu/gazette/story/2012/11/margaret-nast-lewis-101-dies/
11
W H Lewis and M R Lewis, “The Cultivation of Sympathetic Nerves From the Intestine of Chick
Embryos in Saline Solutions,” Anat Record 6 (1912): 7–31.
12
Hematoxylin is a common stain still used in histology. See: https://en.wikipedia.org/wiki/
Haematoxylin
quantity or duration as growths in the plasma media.”13 Tests were made with cells
from liver, heart, intestine, stomach, spleen, adrenal, kidney, limb buds, thyroid,
lung, eye, brain, cord. . . “though not exhaustively.” They suggested that “the
presence of mitotic figures,14 the formation of membranes and the outgrowth of
nerve fibers show that we are dealing with real growth.” Unfortunately, “There are
several types of cells in these outgrowths and we are uncertain as to their identity.
The most common type is probably mesenchymal or connective tissue but even here
uncertainty exists. So that one of the most urgent problems in connection with this
work is the identification of these various outgrowing cells.” In this publication, they
also provided a much more extensive description of the results of the culture of nerve
tissue. Finally, they noted that the addition of a “certain combination of the amino-
acids and polypeptides [to the saline solution, produce] some of the growths [which]
are better than those in simple saline solutions, they are by no means equal to those in
plasma or lymph either in extent or duration.”
The last publication from this initial collaboration examined the types of cell
sheets (termed membranes by the Lewises) that form from specific types of chick
tissues in culture. The most common type, that formed from nearly all the tissues that
were studied, they termed “syncytial” because it was multinucleate, without being
multicellular.15 It was “apparently composed of mesenchymal or connective tissue
cells.” A second “and more interesting” type occurred in cultures of stomach or
intestine and was composed of “small cells closely joined together and may be one or
more layers thick near the old piece but usually spread out towards the periphery to a
single layer of thin flattened cells. . .The edge of the membrane is usually quite
irregular and seems to be amoeboid.” Some cultures, especially those from pieces of
the yolk-sac, displayed a characteristic membrane similar to those from the stomach
or intestine, “except that each cell contains a large clear vacuole or globule close to
the nucleus.” Cultures from heart tissues showed yet another morphology. These
cells were “several times the size of those in the above membrane[s].” The nuclei,
which may be double, were large and “usually [showed] a heavier granule deposit in
the stained preparations.” A fourth “and perhaps most interesting form of membrane
is derived from the kidney tubules.” In this case, the connective tissue cells “wander
away and leave the tubules plainly to be seen” after 24–48 h of culture. Subse-
quently, a “strange irregular membrane. . .spreads out from the broken end of the
tubule” and the cells flattened and irregular and “grow wildly as though some
restraining influence, which normally kept them in order, had been removed.”
They concluded that “it seems that the growth of tissues of the chick embryo
13
W H Lewis and M R Lewis, “The Cultivation of Chick Tissues in Media of Known Chemical
Constitution,” Anat Record 6 (1912): 207–11.
14
Mitosis is the process of cell division that produces two, identical daughter cells. See: https://en.
wikipedia.org/wiki/Mitosis
15
See: https://en.wikipedia.org/wiki/Syncytium
transplanted into artificial media may as in these membranes retain their specific
differentiation and do not return to their embryonic type.”16
In summary, only 2 years into their collaborative research (1910–1912), the
Lewises had demonstrated that they could grow cells from various tissues in a
defined, saline solution, but the growth was not as extensive as that produced
when plasma or lymph was used for the culture medium, and they could not be
sure of the actual cell type that was growing, with the exception of neurons.
Importantly, however, they had also demonstrated that the cultured cells probably
do not de-differentiate back to embryonic cells but retain the differentiation pro-
duced by their previous development.
Margaret also continued her own research during this period. In 1911 and 1912,
she published two papers on aspects of the hematology of tuberculosis and pneu-
monia. She began the 1911 paper by noting that “the fact has long been established
that a cell reacts to its environment and that the condition of any cell at any one time
depends upon its environment.”17 So she wanted to extend this concept to the “state
of the leucocyte18 of the blood and the progress of the disease in tuberculosis.”
Earlier work, in other laboratories, has found that the neutrophils (the most common
subtype of leukocyte) of tubercular patients tended to be largely the immature type
(with simple nuclei), rather than more mature forms (with complex nuclei) found in
healthy humans. These workers had proposed that the distribution of neutrophil
types could be used as a diagnostic feature of tuberculosis. In a typically exhaustive
study of blood cells from a range of patients, Margaret found that the relationship did
not always hold and suggested that a more comprehensive study of all leukocyte
subtypes was necessary for a clear diagnosis of the disease. She concluded that “this
neutrophilic blood picture together with the differential blood count is of great value
in prognosis in tuberculosis, but not of much value in diagnosis.” In a follow-up
study, in collaboration with a Bellevue Hospital physician, James A Miller, Margaret
extended this work to include blood cell morphology from both tubercular and
pneumonia patients, in an effort “to determine their value in the diagnosis and
prognosis of pulmonary tuberculosis and pneumonia.”19 They chose to study the
effects of these two illnesses on blood cell structure to investigate the “contrast of an
acute with a chronic pulmonary disease.” They found that tuberculosis was associ-
ated with an increase in leukocyte number, with an increase in the percentage of
16
M R Lewis and W H Lewis, “Membrane Formations From Tissues Transplanted Into Artificial
Media,” Anat Record 6, no. 5 (1912): 195–205.
17
M R Lewis, “The Blood Picture in Tuberculosis,” Johns Hopkins Hosp Bull 22 (1911): 428–34.
18
The leucocytes, or white blood cells, are part of the vertebrate immune system, which responds to
disease or foreign substances or organisms that enter the body. See https://en.wikipedia.org/wiki/
White_blood_cell. Margaret had intended to study leucocytes for her Ph.D. research at Bryn Mawr;
see Chap. 2.
19
J A Miller and M A Reed, “Studies of the Leukocyte in Pulmonary Tuberculosis and Pneumonia,”
Arch Int Med 9 (1912): 609–36. This paper was based upon blood and patient data sent to Margaret
in 1910, from Dr. Miller; hence, the co-authorship. It is important to remember that Margaret held
only a BA but was now collaborating with a Bellevue MD.
neutrophils, but diminished percentages of the other types of leukocytes (e.g., small
lymphocytes and eosinophils). As earlier works had found, the neutrophils tended to
be more immature than those in the undiseased state. The blood chemistry of
pneumonia patients showed the same leukocytosis, with an increased number of
neutrophils. Patients with a severe case of pneumonia showed the greatest increase in
neutrophils, but the level of immature cells “bears no relationship to the clinical
course of the disease.” These studies by Margaret and her colleagues were surely
some of the earliest to describe the connections between changes in leukocyte
number and subtype percentages with common human diseases. White blood cell
differential counts are now standard measurements when diagnosing human
disease.20
Warren and Margaret returned to the Marine Biological Laboratory at Woods
Hole in the summer of 1912. Interestingly, his academic address was Johns Hopkins,
but Margaret’s address was “1931 East 31st Street, Baltimore,” presumably their
home address—suggesting that she had not obtained a formal academic position at
Johns Hopkins yet,21 although she was apparently an Instructor in Anatomy and
Physiology at Johns Hopkins’ Training School for Nurses in 1911–1912.22 When
they returned to the MBL in the summer of 1914, Margaret’s home institution was
listed as “Johns Hopkins Medical School,” indicating that a more formal association
had been initiated between Margaret and Hopkins.23 Both at the MBL and at their
laboratory at Hopkins, the Lewises had now turned their tissue culture techniques to
an in-depth study of the structure of the newly discovered intracellular organelle
called the mitochondrion.24
As they said in their first paper on this subject (published in the prestigious
journal Science): “The tissues from chick embryos grown outside of the body in
media of known chemical constitution, which we have been studying during the past
3 years, with other problems in mind, have shown such beautiful mitochondria in
both fixed and living specimens that we are led to believe this method offers a better
opportunity for their study than any heretofore used.”25 They used Janus green to
stain “the mitochondria in the living cells a brilliant blue green,” but the color faded
soon (15 min to 3 h). During this short window of time, the cells could be fixed by
placing the coverslip in an osmotic acid vapor from 2 to 5 min. The fixation “is
almost instantaneous and the mitochondria remains practically the same as in living
20
See https://en.wikipedia.org/wiki/White_blood_cell_differential
21
Fifteenth Report of the MBL, 1912, p. 446. For a modern view of the row house, see: https://
www.zillow.com/homedetails/1931-E-31st-St-Baltimore-MD-21218/36462194_zpid/
22
Bryn Mawr College Register of Alumnae and former students, 1920, p. 199.
23
Biological Bulletin, The Marine Biological Laboratory, Vol. 28, No. 6 (1915): p. 351.
24
Although the term mitochondrion had been coined in 1898 by Carl Benda, a variety of other
names had been used during this period, and the origin and function of what we now know as the
“powerhouse of the cell” were relatively unknown in 1912. For a detailed description of the
structure and function of the mitochondrion, see: https://en.wikipedia.org/wiki/Mitochondrion
25
M R Lewis and W H Lewis, “Mitochondria in Tissue Culture,” Science (New York, NY)
39 (1914): 330–33.
cells.” Using this method, the mitochondria could be studied in endothelium,

mesenchyme, giant cells, ectoderm, heart muscle, smooth muscle, and endoderm.
They described that “there are to be found in single stained preparations all grada-
tions of size and shape [of mitochondria] from the small and large granules to short
rods and long rods, to threads of varying length, to anastomosing threads and
networks, which extend throughout the cytoplasm and to rings and loops of various
sizes.” They found, however, that certain cells contained characteristically shaped
mitochondria and that “by far the most important and interesting are the observations
in living cells.” In unstained preparations, the mitochondria were “seen as slightly
refractive opaque bodies. . .[that were] almost never at rest, but are constantly
changing their position and also their shape. The changes in shape are truly remark-
able not only in the great variety of forms, but also in the rapidity with which they
change from one form to another.” But they were unable to determine if mitochon-
dria could divide during cells division, which they wrote “is one of the most
interesting [questions] in the whole field of work upon mitochondria.” They felt
that “the question of the behavior of the mitochondria during the life cycle is one of
great interest and we feel confident that this method of study of the living cell will be
of great value not only for making observations upon the mitochondria, but also for
the study of other activities of the cell.” They concluded this initial publication with
the following statement:
Can we infer from these observations anything concerning the real nature of the mitochon-
dria? Are they organs of the cell, functioning in a definite manner, in other words a living
part of a living cell? If they are organs of the cell, are they concerned in the routine
metabolism which takes place in all living cells or are they concerned with the process of
differentiation of such structure as the myofibrillae, neuro-fibrillae, white fibrous tissue, etc.?
On the other hand, are we dealing with inactive metabolic products of the cell, inactive in the
sense of not being a part of the living protoplasm? If so, are they excretory products which
later are extruded from the cell or storage products which are being continually formed by
the activity of the cell and again used up in its metabolism? What relation do they bear to the
metabolism of the nucleus, if any? The discussion of these most important points must be left
for a more complete account of the mitochondria in tissue culture which is soon to follow.26
This second publication of the Lewises work on mitochondria in tissue culture

was much more extensive—60 pages and 26 figures. And it is clear that some of the
work was done at the MBL because it is listed as their second address, and they
acknowledged the MBL “for the use of a room during the summer of 1914.”26 They
begin by, once again, stating the utility and potential importance of the new tissue
culture protocols:
Tissue cultures afford a new and somewhat different method from that usually employed for
the study of many cell structures. It enables one to compare the living with the fixed material.
In fact, one can study the same cell while living, during the process of fixation, and later as a
stained permanent preparation. It also enables one to follow the changes which take place in
26
M R Lewis and W H Lewis, “Mitochondria (and Other Cytoplasmic Structures) in Tissue
Culture,” Am J Anat 17 (1915): 339–401. It is noteworthy that, one again, Margaret was the first
author of both this and the previous paper in Science.
the living cell from minute to minute. Above all, tissue cultures afford a method by which we
can experiment on the cells and mitochondria. And through such methods only do we
believe a correct interpretation of the significance of mitochondria is to be found.
In spite of the new and different environment of the tissue, i.e., its isolation from the rest
of the embryo; the substitution of a simple Locke’s solution for normal plasma; the contact
with the cover-slip; and the absence of circulation which continually renews the food-supply
and removes the waste, the cells of the tissue cultures are apparently quite normal during the
first two or three days and exhibit no noticeable changes except the characteristic configu-
ration of the growth. How greatly the new environment disturbs the normal metabolic
processes of the cell is impossible to surmise. The cells are in such a thin layer that each
cell is probably as well bathed by the Locke’s solution as in the embryo it would have been
bathed by plasma or lymph. . .The close resemblance of the mitochondria found during this
early period to those found in the chick by other observers. . .shows that they are at least not
noticeably altered in the culture. We are justified, we believe, in assuming that our findings
concerning mitochondria apply as well to the normal cells within the embryo as they do to
the cells of the tissue culture.
Once again, they noted that the living mitochondria “are never quiet, but are
continuously changing in shape, size, and position. . .This extreme plasticity of the
mitochondria is a very important characteristic and was shown in every preparation
examined. It is certainly a feature which must be reckoned with in any attempt to
classify or to analyze their behavior from fixed material.” They found that mito-
chondria could be found throughout the cell, including peripheral extensions, but
sometimes clustered near the nucleus or other cytoplasmic organelles (Fig. 3.2).
They therefore wondered if the arrangement of the mitochondria was governed by
“the shape of the cell, the influence. . .of the nucleus, the internal structure of the
cytoplasm, or do the metabolic activities of the cell govern the size, shape and
arrangement of the mitochondria.” They also studied the number, size, and shape of
mitochondria, all of which varied considerably among adjacent cells in culture. They
summarized these studies by stating that “cells with few mitochondria do not
necessarily have larger mitochondria and there seems to be no definite relation
between size of number. . .The quantity in the cell differs so widely that it has so
far been impossible to connect the quantity of mitochondria with any one factor.
Possibly it is dependent upon the metabolism of the individual cell.”
Previous investigators had wondered if the number of mitochondria increased
during the early stages of mitosis, to provide the mitochondria necessary for the two
daughter cells. The Lewises admitted that “so far, we have only one definite
observation that this is true.” Incubation of one of their cultures at 46 C was
associated with an initial decrease in the size and number of the mitochondria, but
the cells suddenly “began to pass into prophase [the initial stage of mitosis] and
during this process the number of mitochondria in these two cells increased until
they contained more than they had before the experiment was begun.” They con-
cluded that “it seems probable therefore that mitochondria increase in number both
during the resting period and during mitosis; perhaps in some more during the
resting period; in other more during mitosis and in still others during both periods
or only during one.” However, “the daughter cells not only have about one-half the
number of mitochondria found in the mature cells, [and]. . .the mitochondria are
sometimes smaller.” They wondered if the increase in number during the growth
Another random document with
no related content on Scribd:
25
Fritz hat in seiner Klasse zwei Freunde gewonnen, den Kolarczik
Oskar und den Schneider Josef, beide recht üble Bürschchen von
etwas dunkler Abstammung und entschieden mangelhafter
Erziehung. Nach Hause darf er sie nicht mitbringen, denn sie essen
alles mit den Fingern und sind durchaus nicht sauber gewaschen.
Darum hat ihm die Mutter, nach einem ersten probeweisen Besuch,
den Verkehr mit ihnen überhaupt verboten. Natürlich trifft Fritz sie
nun gewissenhaft auf dem Wege von und zur Schule, hat sich in der
Klasse einen Platz in ihrer Nähe gesichert, teilt sein reichliches
Frühstücksbrot mit ihnen, und bringt die kurzen Stunden, die er sich
nachmittag, unter irgendwelchen lügenhaften Vorwänden, von
Hause fortstehlen kann, mit ihnen zu. Das Spielen im Freien ist ihm
nämlich nicht erlaubt, ebensowenig der Besuch von nicht bei ihren
Eltern wohnenden Schulkameraden, und auch diese Besuche nur,
wenn erstens die Eltern anerkannt würdige Leute sind, und zweitens
eine richtige Einladung erfolgt ist. Diese förmlichen
Nachmittagsbesuche aber sind Fritz ein Greuel, denn
unglücklicherweise haben die Leute, die den Eltern als Verkehr
zusagen, meist gräßlich zahme und langweilige Musterkinder. Doch
Widerspruch gibt es nicht. „Deinen Verkehr haben wir zu
bestimmen,“ sagt der Vater. „Ich dulde es nicht, daß du dich mit
Gassenbuben herumtreibst!“
Also gibt Fritz nun fast täglich in den späten Nachmittagsstunden
Übungen des Schülerchors vor, dem er tatsächlich angehört.
Dagegen können die Eltern nichts einwenden, denn der Schülerchor
ist das Steckenpferd des Rektors, und dieser ruft die Jungen wirklich
weit öfter zu Übungen zusammen, als im Unterrichtsplan
vorgesehen ist. Dort steht nur eine Gesangsstunde wöchentlich. Die
leiseste Weigerung der Eltern, ihre Kinder an diesen häufigen
Proben teilnehmen zu lassen, würde der Rektor als persönliche
Beleidigung auffassen. Das alles weiß Fritz genau, weiß auch, daß
die Mutter also nie bei ihren Erkundigungen in der Schule ein Wort
über die vielen Gesangsstunden wagen würde. So lügt er munter los
und weiht die gewonnene Zeit den Freunden. Entweder in dem
weiten, alten Stadtpark, bei wilden und verruchten Spielen, oder,
lieber noch, in dem dumpfigen, düsteren Hinterzimmer, das die
beiden als „Koststudenten“ bei einer ältlichen Witwe zweifelhaften
Gewerbes bewohnen. Das Haus, alt und winkelig, liegt in einem
elenden Seitengäßchen des Armeleuteviertels. Es hat steile, enge
Stiegen, wahre Hühnerleitern, schmale, nie gelüftete Flure, auf die
zahllose Kleinwohnungen münden. Es riecht nach unnennbaren
Speisen, und ewig gibt’s Krawall — Kinder werden geprügelt, Weiber
zanken, oder Eheleute liegen sich in den Haaren. Fritz keucht
jedesmal vor Aufregung, wenn er bei seinen Freunden landet. Wenn
er gesehen und erkannt würde! Der Vater! — Aber die Witwe freut
sich über den feinen Besuch, tätschelt ihm wohlwollend den Kopf
und sagt schmeichelhafte Dinge. Und die Freunde ehren den
Patriziersohn auf ihre Weise, indem sie die Leckerbissen, die er
mitbringt, überschwenglich loben, und gierig verschlingen.
Kolarczik ist übrigens fast fünfzehn, reichlich drei Jahre zu alt für
seine Klasse. Sein Bildungsgang ist etwas verworren, um so reifer
seine Weltanschauung. Er hat Kenntnisse von Wein, Weib und
Gesang, die, im zweiten Punkt, wohl anatomisch falsch oder
unvollständig, jedenfalls aber von keinerlei schamhafter
Zurückhaltung getrübt sind. Er weiß zahllose Liedlein, die mit
kerniger Eindeutigkeit geheime Vorgänge schildern, und singt sie
gern mit seiner brüchigen Wechselstimme. Fritz und Schneider Josef
müssen die Kehrreime mitsingen. Fritz versteht kaum den zehnten
Teil — er ist gänzlich unwissend in geschlechtlichen Dingen, wenn
ihn auch längst schon Ahnungen peinigen. Zu fragen wagt er nicht,
weil er sich keine Blöße geben will. Irgendwann einmal verrät er aber
doch in einer unüberlegten Bemerkung, die gerade recht großspurig
wüst klingen sollte, seine völlige Unschuld. Die Freunde verlachen
ihn furchtbar, lassen sich aber schließlich herbei, ihm auf ihre Art
Bescheid zu sagen. Fritz hört mit heißen Wangen zu. Seine Achtung
vor Vater und Mutter bekommt einen furchtbaren Stoß. — Schöne
Sachen treiben die! Deswegen darf er nie ins Schlafzimmer!
Die Enthüllung beschäftigt ihn nachhaltig und bei irgendeiner
Gelegenheit vermag er der Versuchung nicht zu widerstehen, der
Schwester gegenüber sein Wissen wenigstens anzudeuten. „Was
denn — der Storch! Sei doch nicht so blöd! — Die Kinder werden der
Mutter aus dem Bauch geschnitten, daß du’s weißt!“ Gretl horcht,
wie immer, gläubig und mit regster Anteilnahme. Irgendwelche
Vorstellung verbindet sie mit dem Gehörten nicht.
Kurz darauf kommt Fritz eines Mittags aus der Schule nach
Hause; im Vorraum läuft ihm Gretl arg verweint in den Weg und
macht ihm ein flehendes Warnungszeichen. Doch da ist schon die
Mutter zur Stelle, mit merkwürdig verbissenem Gesicht — wenn sie
die Zungenspitze so zwischen die Zähne klemmt, dann gibt’s was —
faßt den Jungen hart am Arm und führt ihn in die kalte Küche, ganz
am Ende der Wohnung. Dort liegt schon ein Rohrstock bereit. Ein
kurzes Verhör: „Wie darfst du dich unterstehen, der Gretl so unerhört
unanständige Sachen zu erzählen? Und so unverschämte Lügen!?“
— „Aber was ist denn ... ich hab doch nix gesagt ... ich ...“ stottert
Fritz, der wirklich nicht weiß, was los ist. Er denkt an den „Krieg“ —
aber das wär doch nicht so schlimm? — Die Mutter schüttelt ihn:
„Lüg’ nur nicht! Mir machst du nichts vor! — Was sagst du? es nicht
wahr, daß der Storch die Kinder bringt? Und was hast du noch
gesagt?“ — Die Stimme bricht ihr vor Wut und sie beginnt zu
schlagen, lange und mitleidslos. Der dünne Rohrstock zieht
schmerzhafte Striemen über Rücken, Arme und Beine. Fritz heult
furchtbar und denkt dabei doch noch entsetzt, ob wohl „alles“
herausgekommen sei, die Besuche bei den Freunden und das.
Endlich hört die Mutter atemlos auf und fragt ihn abgerissen, böse:
„Wirst du sowas noch einmal sagen? Ha?“ Fritz schüttelt heulend
den Kopf. „Wer bringt die Kinder?“ — Fritz wagt nicht gleich zu
antworten, er wittert eine Falle. Doch da klatscht schon eine böse
Ohrfeige und er stößt fast schreiend heraus: „Der Storch!“ Die Mutter
geht hinaus — er wartet eine Weile, trocknet sich krampfhaft die
Augen und schleicht dann in die Vorderwohnung. Im Kinderzimmer
sitzt Gretl in einem Winkel und weint still vor sich hin. Fritz fühlt
giftigen Haß gegen die Schwester. Er zischt: „Was ist los? Hast du
gepetzt?“ — Gretl verneint entsetzt und weint weiter. Er muß sie erst
mit einem saftigen Puff zum Sprechen bringen. Dann kommt
stockend die Wahrheit: Gretl hat ihren Schulfreundinnen, den
Zwillingen des Apothekers, Fritzens Entdeckung weitererzählt. Und
die Frau Apotheker ist sich zur Mutter beschweren gekommen. ‚So
leid es ihr tue, müsse sie ihren Mäderln wohl den Verkehr verbieten,
denn solche Sachen ...’ Und die Mutter hatte sofort Gretl
vorgenommen, und die mußte wohl oder übel eingestehen, woher ihr
das Wissen gekommen war. Und Gretl hatte, zum ersten Male, einen
Hieb bekommen: „Da schau!“ jammert sie, und zeigt einen blaßroten
Streifen auf dem Unterarm. „Ja, da schau!“ äfft Fritz wütend nach,
streift die Strümpfe von den Beinen und zeigt ihr die
breitaufgeschwollenen dunklen Striemen. „Dumme Gans, mußt du
alles gleich weitertratschen! Ich könnt dich so dreschen!“ — „Und es
ist alles gar nicht wahr!“ trumpft Gretl auf. Doch gleich fällt ihr ein,
was die Mutter noch gesagt hat: „Und die Schande, wenn die
anständigen Kinder gar nicht mehr mit mir verkehren dürfen! Und nur
wegen dir, weil du mich so angelogen hast! Alles gar nicht wahr!“
Fritz kämpft mit dem wütenden Wunsch, den Vorwurf zu widerlegen.
Er weiß genau, daß das mit dem Storch dummer Schwindel ist.
Doch die Selbsterhaltung siegt. Wenn Gretl wieder klatscht, dann
schlagen sie ihn wohl tot. Aber irgendwie muß sich seine Wut Luft
machen. „Ah du, Kohlhobel, verdammter!“ flüstert er. „Seitdem du
auf der Welt bist, ist es nimmer schön für mich! Früher, da war der
Vater gut zu mir, und hat mich in Schutz genommen. Und jetzt krieg
ich Dresch’, und Ohrfeigen, und Schimpfe — alles wegen dir! Bis ich
mich einmal umbring’!“ Und er heult auf, vor Mitleid mit sich selbst.
— Da ist Gretl schon bei ihm, streichelt ihn, spricht ihm zu. „Ich kann
doch nix dafür, schau, Fritzl, ich kann doch nix dafür!“ Und sie weint
herzbrechend. Fritz muß sie schnell beruhigen, denn er hört den
Vater nach Hause kommen. Die Mutter ist bei ihm im Zimmer. Nun
sitzen die Kinder, käseweiß, zitternd, und warten, was noch kommen
soll.
Doch es kommt nichts mehr. Der Vater ist finster, erwidert den
Gruß nicht, spricht bei Tisch kein Wort — aber es geschieht nichts
weiter. Als ob er gar nichts wüßte. Und die Mutter hat es ihm doch
ganz bestimmt gesagt. Und in Fritz regt sich, ganz geheim und
schüchtern vorerst, ein neuer Gedanke: „Aha, er schämt sich — weil
ich die W a h r h e i t weiß! — Aber die Mutter? — Gott, die Mutter —
die kann halt besser lügen!“
Nach Tisch, während die Eltern Siesta halten, schleicht Fritz zur
alten Nanni in die Küche und klagt ihr sein Leid. Er zieht Jacke und
Hemd aus, zeigt ihr den übel zerschlagenen Rücken. Die Alte
stottert vor Erbarmen: „Aber mein Gott im Himmel, so därf man a
Kind doch nie schlagen ... und wegen sowas ... Jesus Maria, ich därf
ja nix sagen ... aber Fritzerle ...!“ Sie weint über ihm, und auch
Fritzens Tränen fließen neu, doch diesmal sind sie süß. Die Alte holt
Läppchen, tränkt sie in Öl, reibt leise, vorsichtig die bösen,
dunkelblauen Striemen ein. Betty, das Stubenmädchen, kommt vom
Aufräumen aus dem Eßzimmer und beteiligt sich an dem
Liebeswerk. Ihre Augen glitzern. Fritz fühlt an seinem bloßen Fleisch
da und dort die Berührung des weichen Mädchenkörpers und ein
ungekanntes Gefühl durchrieselt ihn. Sein Wissen um lasterhafte
Geheimnisse wird rege.
Im Schlafzimmer sagt derweil die Mutter zum Vater: „Ich habe ihn
fest durchgewichst. Das wird er sich wohl merken!“
26
Fritz geht es in der Schule nicht gut. — Nicht, daß er dumm wäre.
Die Lehrer sind sich einig darüber, daß er sogar „einen sehr guten
Kopf“ hat. — Aber sie wissen alle, daß er faul ist, die Hausaufgaben
meist erst in der Schule macht, kurz vor Beginn des Unterrichts, oder
sogar während der Stunde, in der er geprüft werden soll. Manchmal
erwischen sie ihn, meist aber rettet ihn sein gutes Gedächtnis, und
er rutscht an dem „Ungenügend“ vorbei. Auch ist er unaufmerksam,
und schwätzt, und zeigt gelegentlich Ansätze zu Unbotmäßigkeit und
frecher Kritik. Der Ehrgeiz und der verbissene Lerneifer seines
älteren Bruders Felix, rühmlicher Erinnerung, sind ihm fremd.
Darum lieben sie ihn nicht und zwicken ihn gern. Wenn Vater
oder Mutter in die Schule kommen, um nachzufragen, dann erhalten
sie unerfreulichen Bescheid: „Der Junge ist begabt, er könnte viel
leisten. Aber er hat zuviel Allotria im Kopf! Er müßte wohl kürzer
gehalten werden!“ Die Eltern hören den Vorwurf heraus — als ob sie
dem Jungen zuviel Freiheit ließen — und ärgern sich wütend.
Besonders der Vater. Ihm zeigt der Junge bei den eindringlichsten
Strafpredigten, doch auch sonst meist, ein fast dämlich starres
Gesicht, hört wortlos, mit niedergeschlagenen Augen, alles an —
Ermahnungen, Drohungen, Verbote, Befehle. Der Vater fühlt, daß
diese augenscheinliche Fügsamkeit eine Maske ist, hinter der sich
Auflehnung oder Trotz verbergen mögen. Doch scheint ihm der
Gedanke so ungeheuerlich, daß er es vorzieht, den Jungen für einen
Schleicher und Duckmäuser zu halten, für ehrlos kurzum. Das sagt
er ihm auch immer wieder, und Fritzens Liebe für ihn wird dadurch
nicht gesteigert, doch seine Ehrfurcht nimmt Schaden. „Der Alte
kann lang reden,“ sagt Fritz dann verächtlich zur Schwester, „der
weiß viel, wer ich bin!“ Und er erzählt ihr vom Krieg.
Der heimliche Verkehr mit den Freunden dauert an. Fritz hat
ihnen erzählt, daß er wegen der Storchgeschichte Prügel bekommen
hat. Das hat ihren Spott geweckt, aber doch auch eine gewisse
Hochachtung vor dem feinen Haus, in dem die Naturgesetze
willkürlich abgeleugnet werden. Zum Überfluß hat Fritz ihnen
anvertraut, daß er gar nicht der richtige Sohn seiner jetzigen
angeblichen Eltern ist, sondern ein Grafenkind, und daß er bis zu
seinem zehnten Lebensjahr auf einem riesigen Gut, tief drinnen in
Ungarn, aufgewachsen sei, mit Pferden, Hunden und Jagd. „Kannst
du auch Ungarisch?“ fragen sie ihn. „Natürlich,“ sagt er stolz und
spricht einige Sätze eines selbsterfundenen Kauderwelsch. Dann
erzählt er von Bayard, seinem Lieblingsroß, von den Fuchshetzen
und der Jagd auf Bär und Eber, nur mit dem kurzen Spieß und den
treuen Hunden. Er kommt ins Feuer beim Erzählen, und die Freunde
hören ihm gerne zu. Ihr Stillschweigen hat er sich durch furchtbaren
Eid gesichert.
Kolarczik fühlt, daß er sein Ansehen als Ältester wahren muß, da
ihn Fritz sonst ganz in den Schatten stellt. Darum erzählt er von
seinem Vater, der sich totgesoffen habe. „Im Irrenhaus ist er
gestorben. Die Ärzte haben gesagt Desiderium tramons oder
sowas.“ „Desiderium,“ so weit reichten grade noch die
Lateinkenntnisse der drei. Weit entfernt von jeder moralistischen
Ablehnung, empfinden sie vielmehr tiefe Ehrfurcht vor der
merkwürdigen Todesart. Kolarczik hat wieder das Übergewicht. Der
Grafensohn ist beinahe ausgeglichen. Sie möchten selbst zu trinken
anfangen, aber das Geld reicht nicht dazu. Fritz hat zehn Pfennige
Monatsgeld. Sein Vater sagt: „Essen, trinken, schlafen kannst du zu
Hause, Bücher und Kleider werden dir gekauft — also brauchst du
kein Geld!“ Hin und wieder steckt ihm wohl der Herr Rat oder sonst
ein Freund des Hauses ein paar Groschen zu, vielleicht gar eine
Mark. Doch das sind seltene Glücksfälle. Und die Schulbücher kann
man doch nur zu Ende des Jahres verkaufen. Schließlich ergibt sich
ein Ausweg: Schneider Josef gehört zu den armen Schülern, die
vom Hilfsverein für freien Mittagstisch oder monatliche Unterstützung
empfohlen werden. Nun setzt es Fritz bei der Mutter durch, daß er
sich monatlich einen Taler holen darf. Schneider Josef tut natürlich
begeistert mit; so oft er aber seine Spende einkassiert hat, lauert ihn
Fritz auf der Treppe ab und läßt sich sein Drittel, eine Mark,
auszahlen. Dann geht es auf der finsteren Studentenbude ein paar
Tage lang hoch her, mit Bier, Rollmöpsen und Zigaretten. Wein ist zu
teuer, und Schnaps ist zu scharf. „Aber von Bier muß man so viel
trinken, bis man richtig besoffen ist,“ meint Kolarczik. „Das ist fad! —
Ihr habt’s doch so einen großen Weinkeller — schau halt, daß du ein
paar Flaschen klauen kannst!“ Fritz wehrt sich zunächst — die
Gefahren des Unternehmens sind ungeheuer. Aber die Freunde
lassen nicht nach: „Ein schöner Grafensohn!“ höhnen sie. „Wenn’s
einmal heißt, Kurasche haben, macht er in die Hosen!“ Das gibt den
Ausschlag. Kurz darauf bringt Fritz zwei dickverstaubte Flaschen an.
Mehr hat er nicht unbemerkt fortbringen können. Es ist uralter
Bordeaux, ganz trüb vom Schütteln. Sie trinken ihn aus
schlechtgespülten Biergläsern und singen wüste Lieder dazu. Die
Witwe kommt herein und tut mit. Sie setzt sich neben Kolarczik, der
sie dreist anfaßt, schlenkert mit den Beinen, daß man die groben,
blau und weiß geringelten Strümpfe bis zum Knie hinan sieht. Sie ist
recht unsauber und riecht stark. Fritz ist dabei übel zumut. Er fühlt
sich als todeswürdiger Verbrecher, hat entsetzliche Angst vor
Entdeckung — und ist doch prickelnd aufgeregt und stolz. — Bevor
er nach Hause geht, rennt er im bloßen Rock in der Winterluft auf
und ab und kaut Pfefferminz, um den Tabak- und Weingeruch
loszuwerden. In der Dienstbotenkammer zieht er rasch den
Hausanzug an und versteckt die verrauchten Kleider. Betty ist ihm
beim Umziehen behilflich, mit merkwürdig eindringlichen Griffen.
Fritz hält sich zeitweilig an ihrer Brust an, und sie läßt es geschehen.
„Die ist verrückt auf mich!“ denkt er. Aber sein Sieg macht ihm Angst.
27
Kolarczik und Schneider haben, trotz Eid, das Geheimnis von
Fritzens hochadliger Abstammung nicht gewahrt. — Bei der Witwe
erscheint des öfteren ein Schlossergesell, begeisterter
Gelegenheitsarbeiter und Brotzeitmacher, im Nebenberuf wüster
Lebemann. Den haben die beiden ins Vertrauen gezogen, er aber
hat die Enthüllung mit brüllendem Hohngelächter aufgenommen. Er
kennt, wie die ganze kleine Stadt, den Sanitätsrat und seine Familie
genau, kann sich an Fritzens und seiner Brüder Geburt und
Kinderjahre genau erinnern und läßt keine Silbe von Fritzens kühner
Dichtung bestehen. Letzte Zweifel der Buben zerstreut er, indem er
seinerseits die Witwe als Zeugin anruft. Die stimmt ihm natürlich bei
und verspottet die Jungen wegen ihrer unerhörten Leichtgläubigkeit:
„Na, ihr seid’s mir schöne Eseln! Wenn ihr auch vom Dorf
hereingekommen seid’s — aber s o blöd braucht’s ihr euch doch
nicht anschmieren lassen! — D e r ein Sohn von einem Grafen! —
Ich weiß noch wie heut — die Frau Czepinka, die Hebamme, hat ihn
gebracht — damals haben sie am untern Markt gewohnt, in dem
großen Haus, wo unten der Sattler Malik die Werkstatt hat! Da
schaut man’s wieder: die Kinder von die feinen Leute! S o lügen! Ob
ich mich nie schämen möcht’!“
Gleich am nächsten Tage fielen sie über Fritz her und
überschütteten ihn mit hohnvollen Vorwürfen so beißend und
unerschöpflich, daß er gar nicht dazu kam, sie wegen des
gebrochenen Eides zur Rede zu stellen. Kolarczik schnitt ihm gleich
das Wort ab: „Du mit deinem Eid! Du kannst mir am Buckel steigen!
So a Liegner! Weißt du, was du bist? Du bist a großer Schöps! Das
bist du!“
So hatte die Freundschaft ein jähes Ende. Furchtbare Drohungen
wurden ausgestoßen, doch blieb es dabei. Sie wußten gegenseitig
zuviel „auf einander“ und keiner konnte wirksam den andern
verpetzen, ohne selbst mit hineinzufallen. Eine kleine Genugtuung
für Fritz war es, daß er die Einstellung der monatlichen
Unterstützung an Schneider Josef durchsetzte. Zuerst hatte
Kolarczik wohl gedroht, Fritzens Eltern „alles zu sagen“. Aber Fritz
hatte ihm wütend entgegengehalten, daß erstens der Sanitätsrat
einen so schmierigen Jungen wie den Kolarczik gar nicht einmal
anhören würde. Und wenn schon, und wenn er, Fritz, auch
totgeschlagen würde — vorher würde auch e r noch „alles sagen“ —
von der Kostbude, und dem Verhältnis zu der Witwe, und alles
überhaupt — und dann würden sie alle aus der Schule
hinausgeschmissen, und die Witwe würde sicher eingesperrt. Da
hatten sich Kolarczik und Schneider zähneknirschend gefügt. Denn
des einen Mutter wie des andern Vater befaßten sich zwar selten mit
der Erziehung ihrer Sprößlinge, dann aber höchst gründlich, und
schlugen beide keine schlechte Klinge. Und Fritzens Rachsucht
kannten sie. Er war „ein Luder“!
Noch eine Folge hatte das Ereignis: als Gretl wieder einmal vom
„Krieg“ hören wollte, sagte ihr der Bruder mit dürren Worten, daß er
all die Geschichten nur erfunden habe. Gretl stürzte aus allen
Himmeln — Lüge war ihr überhaupt unfaßbar — und nun eine so
ausgedehnte, furchtbare, durch Jahre hingesponnene Kette von
Lügen! Die blinde Verehrung, die sie dem Bruder für seine
vermeintlichen Heldentaten gezollt — die rasende
Selbstüberwindung, die sie aufgebracht hatte, um das Geheimnis zu
wahren — die Gebete vor dem Schlafengehen, der Bruder möchte
bei dem heimlichen Verlassen des Hauses oder morgens bei der
Rückkehr nicht erwischt werden —: alles umsonst! Sie weinte
fassungslos. Es war die erste große Enttäuschung.
Wenn ihn auch die Ablehnung der Schwester, die gehässige
Verachtung der ehemaligen Freunde schmerzten, so fühlte Fritz
doch starke Erleichterung bei dem Gedanken, daß er sich hinfort
nicht mehr anzustrengen brauchte, die einmal erfundenen Lügen
krampfhaft durchzuhalten. Und fein war es doch! Der Schneider
Josef war ja ein Trottel — aber daß Kolarczik, der alte Gauner, ihm
so lange geglaubt hatte — das war schon was wert! — Wegen Gretl
schlug ihm wohl das Gewissen — sie hatte so blind vertrauensselig
geglaubt. Und war nun so furchtbar traurig. Das sah er nicht gern.
Doch hatte diese Entfremdung auch ihr Gutes: Seit der
Storchgeschichte war er fest entschlossen, der Schwester nie wieder
von gewissen Dingen zu reden. Und wieviel Geheimnisse hatte er
nun zu hüten: Das Rauchen, Trinken, die gelegentlichen Griffe in
den Weinkeller, sein reiches Wissen um „die Weiber“, und vor allem:
B e t t y . Mit der war er zu einem merkwürdigen Zustand gekommen:
wo immer er sie erwischte, fing er mit ihr zu raufen an, und sie hielt
ihm meist leidenschaftlich Widerpart. Er boxte sie heftig, rang mit ihr,
warf sie zu Boden oder auf ein Sofa — sie ließ ihn mit heißen Augen
gewähren, preßte ihn an sich, zog ihn mit in ihren Sturz. Dann fühlte
er durch das dünne Kleid die weiche warme Mädchenbrust und
glaubte zu vergehen in dunkler Sehnsucht. Oft endeten diese
Zwischenfälle eigenartig — das Mädel machte sich plötzlich aus
seiner Umschließung frei, wehrte ihn derb ab, wenn er sie halten
wollte, blitzte ihn verächtlich an und lief davon. Einmal nannte sie ihn
sogar einen dummen Jungen. Fritz blieb gedemütigt, ratlos zurück
und brütete Rache.
Abends beschwerte er sich bei der Mutter, daß seine Schuhe und
Kleider so schlecht geputzt würden. Und Betty bekam einen
scharfen Verweis.
28
Weitaus das langweiligste aller Schulfächer war der katholische
Religionsunterricht. Doch man durfte sich nichts merken lassen,
denn der Katechet war aus Härte und Milde unberechenbar
gemischt, und wen er einmal „auf dem Strich“ hatte, der durfte sich
vorsehen. Fritz war ohnedies nicht sein Liebling, denn Felix war
während seiner letzten Schuljahre in dem Verdacht gestanden, daß
er „nichts glaube“ — und das Mißtrauen hatte sich auf den jüngeren
Bruder übertragen. Gegen Menschen aber, von denen er fühlte, daß
sie ihn nicht bedingungslos liebten, empfand Fritz sofort Haß, der, je
nachdem, mit Furcht oder Rachgier gemengt war. Den Katecheten
also haßte er leidenschaftlich — doch was konnte er ihm zum Trotz
tun? Der Katechet war nahezu allmächtig in der Lehrerkonferenz;
der Rektor war streng klerikal.
Die Klasse hielt bei der katholischen Liturgie. Der Katechet
verbreitete sich salbungsvoll und eindringlich über das umständliche
Zeremoniell einer Bischofsweihe — wieviel assistierende Priester,
wieviel Ministranten, wieviel Rauchfässer, und in welcher
Reihenfolge sie geschwungen werden müßten. Fritz belauerte jedes
seiner Worte, um einen Anlaß zu geheimem Spott zu erhaschen,
einen Zungenfehler etwa, oder eine ungewollte Zweideutigkeit. Doch
der Katechet versprach sich nicht, blieb nicht stecken — ölig und
ungehemmt strömte sein Vortrag. Mit einem Mal schoß Fritz ein
Gedanke durch den Kopf, so heftig und unvermittelt, daß er fast
erschrak: „Das ist lauter Blödsinn, das Räuchern und Läuten —
davon hat der Herr Christus nix gewußt!“ Entsetzt starrte er den
Katecheten an, ob der ihm etwa die lästerlichen Gedanken von der
Stirn ablesen würde. Doch nichts geschah. Die Stunde ging weiter.
Beim Schlußgebet faltete Fritz zwar die Hände und bewegte die
Lippen, die Worte aber sprach er nicht mit. Er summte nur den
leiernden Tonfall der Klasse nach.
Abends ließ er, zum erstenmal und nicht ohne Angst, das
Nachtgebet ungesprochen. Lange wälzte er sich schlaflos, kämpfte
mit dem Wunsch, doch noch, im Dunkel, unter der Decke, die Hände
zu falten und das Versäumte nachzuholen. Der Gedanke aber: wie
wütend der Katechet sich ärgern würde, wüßte er, daß Fritz nun
auch „nichts glaube“ — der Gedanke ließ ihn stark bleiben. Im
Hochgefühl befriedigter Rache schlief er endlich ein.
In einer der nächsten Stunden kam der Katechet auf die
unsterbliche Seele zu sprechen, die Alleingut des Menschen sei, und
ließ sich abschweifend über die Unvernunft des Tieres aus. In Fritz
hatte der Unglaube inzwischen kräftig Wurzel geschlagen und er
verspottete insgeheim die kurzsichtige Verblendung des Lehrers. Zu
Hause erwartete ihn, freudig bewegt wie immer, Flocki, ein
schwarzer Zwergspitz, der, wie die meisten Familienhunde, bis zu
hohem Grade zum Verständnis der menschlichen Gewohnheiten
und Worte vorgedrungen war. Fritz stürzte auf ihn zu, mit
stürmischen Liebkosungen, balgte, hetzte mit ihm, kniete endlich zu
ihm nieder und fragte leidenschaftlich: „Flocki! — sie sagen, du bist
unvernünftig! Flocki! Solche Esel! Was? Flocki!“ — Der Hund saß vor
ihm und sah ihm starr ins Gesicht. Die Worte waren ihm fremd, und
er suchte zu ergründen, was man von ihm erwarte. Der Junge
wiederholte seine Frage, wilder, eindringlicher. Flocki verstand, daß
man eine Gefühlskundgebung wünsche, bellte kurz auf und fuhr dem
Jungen mit der Zunge ins Gesicht. Es war tatsächlich die beste
Antwort, die er geben konnte. Fritz war begeistert, und da eben
nichts Besonderes gegen ihn vorlag, wagte er sogar bei Tisch davon
zu erzählen. Er sah genau, wie der Vater schmunzelte, und fügte,
kühn gemacht, noch die Andeutung sonstiger Zweifel hinzu. Keine
Zurechtweisung kam. Fritz war namenlos stolz. Nach Tisch aber rief
ihn der Vater ins Arbeitszimmer. Da wich der Stolz rasch arger
Beklommenheit — vielleicht kamen jetzt die Prügel nach? Man
konnte nie wissen! Doch der Vater war nicht böse, sprach ganz
freundlich: „Ich kann dich nicht tadeln, wenn du dir eigene Gedanken
machst — du wirst ja allmählich ein großer Junge! Aber das bitte ich
mir aus, daß du das für dich behältst, in der Schule natürlich
sowieso, vor allem aber der Gretl gegenüber! Weh’ dir, wenn ich
einmal erfahre, daß du dem Mädel etwa Grillen in den Kopf setzest!
Merk’ dir das! Und jetzt geh!“ Und Fritz ging beseligt. Die Mutter kam
ihm ins Kinderzimmer nach und sagte mit nassen Augen — sie war
leicht gerührt —: „Da siehst du wieder, was du für einen guten Vater
hast!“ Dann ließ sie ihn allein. Fritz wartete, bis sie aus dem Zimmer
war, und äffte ihr stumm nach: „Ja, guter Vater! — Weil er mich nicht
gedroschen hat, wie du wegen dem Storch! Und ihr glaubt’s ja alle
beide selber nix! — Der Alte geht überhaupt nie einmal in die
Kirche!“ — Und er kam sich groß und gereift vor. Dann schlich er ins
Speisezimmer, wo Betty den Tisch abräumte; das Mädchen war ihm
noch böse wegen seiner letzten Angeberei. Doch er brach ihren
Widerstand, faßte sie kühn, wie nie zuvor, zwang sie auf die breite
Ottomane nieder. Da kam plötzlich ein neuer Ausdruck in ihr
Gesicht, sie wurde weich, widerstandslos unter ihm, ihre Schenkel
wichen. — Er sprang auf und rannte atemlos davon.
29
Kurz darauf kam die Nachricht nach Hause, Fritz sei von der
Lehrerkonferenz in Sitten getadelt worden, wegen Störung des
Religionsunterrichts durch Schwätzen und Allotria. Mit diesem
Sittentadel hatte es seine eigene Bewandtnis: Fritz hatte, bei der
Erörterung der himmlischen Rangordnung, eine der Form nach
einwandfreie Zwischenfrage getan — ob nämlich die Engel ewig in
ihren Dienststufen verbleiben müßten, oder ob sie auch befördert
würden? Aber in Tonfall oder Mienenspiel hatte er wohl den richtigen
Ernst allzu unvorsichtig vermissen lassen. Und überdies wußte der
Katechet im Augenblick tatsächlich keine Antwort auf die
unerwartete Spitzfindigkeit. So goß er erst die Schale seines
kanonischen Zornes über den frechen Frager aus und behielt ihn
dann scharf im Auge. Und als Fritz, der das Unwetter vorbei wähnte,
endlich grinsend einen beifallsuchenden Rundblick durch die Klasse
schickte, da griff der Katechet zu, trug ihn ins Klassenbuch ein und
verschaffte ihm den Sittentadel. Es war ein offensichtlicher
Racheakt, wohl auch eine kleine Rechtsbeugung. Doch angesichts
des Tadelbriefes ließen die Eltern keinerlei Rechtfertigung zu, hörten
die Darstellung des Jungen gar nicht an. Der Vater prügelte ihn
furchtbar, die Mutter aber bestand überdies noch darauf, er müsse
mit ihr zum Katecheten gehen und um Verzeihung bitten. Das war
das Schlimmste, weit ärger noch als die Prügel, obwohl ihm davon
noch jeder Knochen im Leibe weh tat. Er bat flehentlich, ihm den
Bittgang zu erlassen. Ihm wurde fast übel bei dem Gedanken, den
tödlich gehaßten Schwarzrock um Verzeihung bitten zu sollen. Doch
die Mutter gab nicht nach, faßte ihn am Arm und führte ihn, ganz
verweint und verschwollen im Gesicht, mit sich. Im Vorflur des
großen alten Stiftshauses, in dem der Katechet wohnte, machte Fritz
einen letzten verzweifelten Versuch, der furchtbaren Demütigung zu
entrinnen. Da wurde die Mutter böse, schlug ihn heftig ins Gesicht,
krallte ihm die Finger in den Oberarm — sie kannte die
empfindlichen Stellen — und zog die Glocke. Ein hallender Erzton
durchzog das weite Haus, dann öffnete ein sanfter Pförtner die Tür,
fragte nach dem Begehr der Besucher und führte sie durch lange
Gänge, bis vor eine schmale Bogentür. Dort klopfte er an und ging.
Die Tür wurde von innen aufgetan, und der Katechet stand auf
der Schwelle. In seinem Gesicht kämpfte Genugtuung und
Rachsucht, als er Mutter und Sohn erblickte. Er bat sie ins Zimmer,
und die Mutter setzte ihm auseinander, wie sehr sie und ihr Mann
über das Benehmen des Jungen empört seien; er habe strenge
Strafe bekommen und nun habe sie ihn hergebracht ... Der Katechet
unterbrach sie höflich und salbungsvoll, sagte einiges, was Fritz
nicht verstand. Haß und das Gefühl völliger Ohnmacht erstickten
den Jungen fast. Da hörte er die Mutter sagen: „Also vorwärts —
bitte um Verzeihung!“ Und wieder zwickte sie ihn unmerklich in den
Arm. „Ich ... ich ...“ stotterte Fritz schluchzend. Immer noch hoffte er,
das Aussprechen der widerlichen Formel werde ihm erspart bleiben.
Doch der Katechet sagte gedehnt: „Nun ...?“ und Fritz hörte deutlich
den beißenden Hohn des Siegers. Der schenkte ihm nichts! Da
gruben sich abermals die Finger der Mutter in seinen Arm, und er
stieß hervor: „Ich ... bitte ... um ...“ — das Wort „Verzeihung“ aber
ging in furchtbarem Aufheulen unter. Der Katechet wollte wohl den
Bogen nicht überspannen. Er gab sich zufrieden, gab süß lächelnd
der Mutter einige Ratschläge, die ebenso viele giftige Spitzen waren
und strich Fritz mit der Gebärde der Vergebung über den Kopf. Dem
Jungen rann die Berührung der weichen, fetten Priesterfinger wie
Gift durch den Körper. Dann zog ihn die Mutter hinaus.
Von da an zog Fritz nicht mehr den Hut, wenn er an einer Kirche,
einem Kruzifix oder Heiligenbild vorbeikam, und zischte unflätige
Lästerungen durch die Zähne: „Krepierter Hund!“ — Das war seine
Rache.
30
Kolarczik und Schneider haben sich an Fritz herangemacht und
mit Honigmienen Versöhnung angeboten. Sie quellen über von
reumütigen Erinnerungen an die schöne Zeit von ehemals. Fritz
spielt zunächst den Unerbittlichen, doch schmeichelt ihm die
Unterwürfigkeit der andern doch so sehr, daß seine Widerstände
rasch dahinschmelzen.
Im Stadtpark hausen die „Parkindianer“, das sind Kinder der
untersten Schichten, von rohen Sitten, Schüler der Volks- und
Fortbildungsschulen; auch viele Handwerker — und Fabrikslehrlinge
sind darunter. Sie nennen die Gymnasiasten „Gimpel“ und die
Realschüler „Retiradschüler“. Denn sie hassen alles, was eine
höhere Schule besucht oder einen sauberen Rock trägt. Sie sind ein
unzuverlässiges Volk, halten keine Bündnisse und kehren sich nicht
an ritterliche Kampfregeln. Sie greifen nur an, wenn sie in der
Überzahl sind, dann aber rücksichtslos und ohne Anlaß. „Gelbmütz“
und „Rotkappel“ sind ihre Führer, gefürchtet wegen ihrer Roheit. „Die
schmeißen mit Dreck,“ behauptet die Sage.
Für Fritz und seine Freunde ist ein Gang in den Stadtpark stets
mit der Gefahr verbunden, von einer erdrückenden Übermacht
verprügelt zu werden. Denn wenn auch nur ein einzelner
Parkindianer sie erblickt, dann ruft er mit gellendem Pfiff aus den
Arbeiterhütten am Parkrand Verstärkung herbei und die Hetzjagd
beginnt.
Fritz und seine Freunde g e h e n nicht — sie geben stets vor,
mutige Renner zu reiten. Beim Verlassen der Stadt reißt sich jeder
eine Gerte ab, die als Reitpeitsche dient. Damit klopfen sie sich die
Waden und tänzeln dahin, während die steifgehaltene linke Faust
gedachte Zügel handhabt. Dringt Übermacht auf sie ein, so rasen sie
davon und geißeln sich wütend. Fritz besonders kann gänzlich das
Gefühl verlieren, daß die Beine zu ihm gehören. Er schlägt sich
blutige Striemen, während er atemlos vor den Verfolgern dahinjagt.
Noch nie ist er eingeholt worden. Kolarczik und Schneider können
auch gut rennen, doch ist er ihnen immer meterweit voraus. Sind sie
dann in Sicherheit, so klopft er stolz und dankbar seine schlanken
Schenkel. Er hat sie „Falko“ getauft.
An einem Winternachmittag ziehen die frisch versöhnten Freunde
hinaus in den Park. Fritz tänzelt und kurbettiert auf Falko.
Weit draußen, am Ufer des dick gefrorenen Flüßchens, finden sie
ein halbes Dutzend Parkindianer dabei, auf dem Eis eine lange
Rutschbahn herzustellen. Es sind lauter kleine Burschen, die trotz
ihrer Überzahl keinen Angriff wagen. Die drei Stadtbuben wollen
aber auch nicht anfangen, denn sie wissen, daß ein kleiner
Anfangserfolg sich unweigerlich in eine schließliche Niederlage
verkehren müßte. Die herbeigepfiffene Verstärkung würde ihnen den
Rückweg durch den Park verlegen. — So kommt es zu einem
friedlichen Einvernehmen, die Parteien bemühen sich einträchtig um
die Rutschbahn. Die ist unheimlich glatt und lang und endet wenige
Meter vor einer offenen Stelle, wo das schwarze wirbelnde Wasser
durch das Eis dringt. Es handelt sich darum, richtig Anlauf zu
nehmen, die Bahn stehend zu durchfahren und möglichst knapp vor
dem Loch im Eis mit kurzem Schwung anzuhalten. Das Spiel ist
fesselnd. Fritz treibt es toller als die andern. Mit wahrer Wollust läßt
er sich bis hart an die Kuhle gleiten. „Weil der Vater immer sagt, ich
bin ein Feigling!“ denkt er mit zusammengebissenen Zähnen. „Der
weiß viel!“
Da dröhnen durch die dunstige Winterluft die Schläge der
Stadtuhr herüber: Dreiviertel vier! Und um vier ist Gesangsstunde,
diesmal aber wirkliche Gesangsstunde. Im Laufschritt könnte man
noch annähernd zurechtkommen, die kleine Verspätung wäre leicht
zu entschuldigen. — Während Fritz noch überlegt, treten die beiden
Freunde auf ihn zu: „Du denkst wegen der Gesangsstunde? Sei
doch nicht blöd — jetzt wird’s grad schön, da werden wir doch nicht
schon weggehen! Schwänzen wir halt! Es kommt ja nicht raus! Und
singen können wir hier überhaupt auch!“ Und Kolarczik stimmt ein
kerniges Lied an. Fritz ist schnell verführt. Und sie tollen weiter auf
dem Eise. Knapp nach fünf brechen sie auf, durchjagen den
nächtigen Park — hinter jedem der bereiften Büsche können Feinde
lauern — und zu angemessener Zeit landet Fritz im Elternhaus.
Falko ist prachtvoll galoppiert, allerdings nicht ohne Peitsche.
Am nächsten Morgen wird Fritz in der Klasse mit den entsetzten
Gesichtern, den Gebärden des Abscheus empfangen, wie sie die
Mitschüler den überführten Verbrechern aus ihren Reihen zu zeigen
pflegen. Er verlangt Aufklärung, doch alle ziehen sich scheu vor ihm
zurück, besonders heftig Kolarczik und Schneider. Die erste Stunde,
Mathematik, hat der Klassenlehrer. Er ruft sofort Kolarczik,
Schneider und Fritz zu sich aufs Katheder und beginnt das Verhör:
„Warum habt ihr gestern die Gesangsstunde geschwänzt?“ Und
bevor Fritz noch den Mund auftun kann, um die verabredete
Entschuldigung vorzubringen, legt Kolarczik schon heulend los. —
Es ist schmählicher Verrat. — Aus Rache! Fritz ist in eine plumpe
Falle geraten. Er steht starr vor Entsetzen über soviel Niedertracht:
Früh morgens schon waren Kolarczik und Schneider zum Direktor
gerannt und hatten weinend gebeichtet, sie hätten sich von Fritz
verleiten lassen, die Gesangsstunde zu schwänzen und sie wollten
es auch nie, nie, nie wieder tun. Und man möchte sie doch um
Gottes willen nicht bestrafen, ihre Kostfrau sei so streng, und ihre
Eltern noch mehr. Und Fritz sei doch allein an allem schuld. — Ihr
Fernbleiben war tatsächlich gar nicht bemerkt worden. Und da sie
trotzdem — dies unterstrich der Klassenlehrer salbungsvoll —
freiwillig ein reumütiges Geständnis abgelegt hätten, so sei
gnadenweise von einer Bestrafung abgesehen worden. Nicht so bei
Fritz, dem böswilligen Anstifter. „Du wirst vielleicht“ — die Stimme
des Lehrers knarrte vor Verachtung — „durch gehäufte Lügen deine
Schuld zu mindern, dich der wohlverdienten Strafe zu entziehen
trachten! Versuche es nicht, zur Schuld noch die Schande zu fügen!
Du bist überführt durch das gleichlautende Zeugnis deiner armen
Mitschüler! Schweige also!“ — Doch Fritz dachte gar nicht an
Verteidigung. Er war wie gelähmt und nahm wortlos das harte Urteil
hin: Zwei Stunden Karzer und Bestätigung vom Vater. — Als er
stumm auf seinen Platz zurückging, knarrte ihm die böse Stimme
nach: „Ein verstockter Sünder! Nun, ich hoffe, daß dir dein Vater das
Verständnis für die Tragweite deiner Handlungsweise erschließen
wird! Haha!“ Die Klasse gröhlte pflichtschuldig Beifall. Kolarczik aber
warf ihm hinter vorgehaltenem Buch einen Blick voll teuflischer
Schadenfreude zu.
Und zufällig war am selben Vormittag die Mutter in der Schule
und hörte vom Direktor brühwarm die empörende Nachricht. Sie
empfing den Knaben zu Hause, bleich und bebend vor Entrüstung,
sprach nur wenige schneidende Worte und schickte ihn in die kalte
Küche, um die Heimkehr des Vaters abzuwarten. „Ich schlage dich
nicht — o nein! Das kann der Papa besser! Warte nur, bis er kommt!“
Der Vater kam gerade diesmal mit arger Verspätung, müde und
hungrig, und war doppelt wütend, daß er noch mit Erziehungsfragen
aufgehalten wurde. Er fand den Buben halb ohnmächtig von dem
stundenlangen Warten auf die Exekution. Irgendeine Verteidigung
ließ der Vater so wenig zu, wie es die Mutter getan hatte. Der Fall
lag ja sonnenklar. Und die Züchtigung fiel darnach aus. Mehr noch
als die gewiß vollwichtigen Prügel schmerzte den Jungen aber die
furchtbare Beschimpfung: „Ehrloser Schuft!“ — Und doch erfüllte ihn
das Bewußtsein, daß er alle die Qualen zu Unrecht erduldete, mit
grausamer Freude. Er zog nicht einmal Gretl nachher ins Vertrauen,
auch Nanni und Betty nicht. Dieses stumme Leiden tröstete ihn.

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Full Ebook of Pioneers in Cell Physiology The Story of Warren and Margaret Lewis 1St Edition David H Evans Online PDF All Chapter

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Pioneers in Cell Physiology The Story

of Warren and Margaret Lewis 1st

Mechanisms in Cell Physiology Michele Mazzanti

Brain Physiology and Psychology C. R. Evans (Editor)

Soil The Incredible Story of What Keeps the Earth and

The Human Gene Editing Debate 1st Edition John H. Evans

Big Road Blues David Evans

Fear of the Father The Crockett Series 1 1st Edition

Scientific Debates in Space Science Discoveries in the

Hoosiers and the American Story 1st Edition James H.

Published on behalf of The American Physiological Society

Pioneers in Cell Physiology:

ISSN 2625-2813 ISSN 2625-2821 (electronic)

Jointly published with The American Physiological Society

Gainesville, FL, USA David H. Evans

I am delighted to have the opportunity of sending. . .a message of warmest greetings to

1 Warren Lewis: Early Life, Educational Background, and Early

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 1

artiﬁcial parthenogenesis, and heteromorphosis (“the replacement of an injured or removed organ

constitution of living matter.” Since potassium cyanide was known to inhibit “a

epidermis. These experiments suggested that lens development was dependent on

In a subsequent publication (also published in 1904), Warren gave a much more

Warren extended his studies of eye development in a paper, published in 1907,

Fig. 1.5 Copy of a

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 19

developmental biologist and geneticist.14 Margaret remained at Bryn Mawr for

(1906–1908) of Johns Hopkins’ Ross Harrison on the mechanisms of amphibian

Margaret and Warren were married on May 23rd, 1910.

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 29

Fig. 3.1 Photomicrograph

cells.” Using this method, the mitochondria could be studied in endothelium,

This second publication of the Lewises work on mitochondria in tissue culture

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