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Joseph Needham, F.R.S.
Joseph Needham, F.R.S.


Fellow of Grmville E‘! Cain: College, and Sir William Dunn Reader in Biochtnrimy
Fellow of Grmville E‘! Cain: College, and Sir William Dunn Reader in Biochemisty


in the University of Cambridge
in the University of Cambridge


Second Edition Revised With The Assistance Of
Second Edition Revised with the Assistance of


Arthur Hughes, Ph.D
Arthur Hughes, Ph.D


Letlmer x'nAmztamy in the University of Cambridge
Lecturer in Anatomy in the University of Cambridge


CAMBRIDGE
Cambridge
AT THE UNIVERSITY PRESS
 
At The University Press


1959
1959
PUBLISHED BY
Published By
THE SYNDXC3 OF THE CAMBRIDGE UNIVERSITY PRZS5
The Synod Of The Cambridge University Press, London 0559: Benllty House
London 0559: Benllty House




Line 38: Line 38:




CONTENTS
==Contents==
 
Preliminary Note
List of Plates 9
Preliminary Note 1 r
Chapter One
EMBRYOLOGY IN ANTIQUITY
1. Ideas of Primitive Peoples 18
2. Egyptian Antiquity 18
3. Artificial Incubation 22
4. Indian Antiquity 25
5. Hellenic Antiquity; the Pre-Socratic: 27
6. Hippocratic Embryology and the Doctrine of the Two Seeds 3!
7. Aristotle’s great Systematisation 37
8. The Doctrine of the Menstrual Blood 42
9. Denials of Maternity and Paternity 43
lo. Formation, Recapitulation and Fermentation 46
H. The Aristotelian Bala.noe—sl1eet 54.
I2. AristotIe’s Theory of Causation 56
13. The Hellenistic Age 60
14. Galen and the Vital Faculties 69
 
Chapter Two
EMBRYOLOGY FROM GALEN TO THE RENAISSANCE
1. Patristic Speculation 75
2. Contributions of Jewish Thinkers 77
3. Embryology among the Arabs 82
4. Alchemy and Embryology 83
5. The Visions of St Hildegard 84
6. Albertus Magnus; the Re-awakening of Scientific Embryology 86
7. Aristotle’: lllaxterpiece 91
8. Scholastic Ideas on Generation 93
9. The Insights of Leonardo da Vinci 96
10. The Macro-Ioonogrnphers of the Sixteenth Century 99
II. The Movement to Rationalise Obstetrim 109


Chapter Three
List of Plates


EMBRYOLOGY IN THE SEVENTEENTH CENTURY
[[Book - A History of Embryology 1959-1|'''Chapter One - Embryology in Antiquity''']]
1. The Opening Years "5
# Ideas of Primitive Peoples
2. Developmental Deter-rninism and Tmnsplantation; Digby, Higlunom and Tagliacozzi my
# Egyptian Antiquity
3. Thomas Browne and the Beginnings of Chemical Embryology ,3,
# Artificial Incubation
4. Willizun Harvey and the Identification of the Blastoderrn 133
# Indian Antiquity
5. The Riddle of Fertilisation 145
# Hellenic Antiquity; the Pre-Socratic
6. Harvey's Achievemcms and Xnfluenu: I49
# Hippocratic Embryology and the Doctrine of the Two Seeds
7. Atomist Theories of Embryonic Development; Gaucndi and Dcscm-ta I53
# Aristotle’s great Systematisation
8. Fixatives and Uterine Milk; Robert Boyle and Walter Needhzun r 58
# The Doctrine of the Menstrual Blood
9. The Discovery of the Follicles of the Mammalian Ovary 162
# Denials of Maternity and Paternity
10. The Miuro-iccnogr-aphers and Preformatianism; Marcello Malpighi and Jan Swammerdam 163
# Formation, Recapitulation and Fermentation
11. Foetal Respiration and Composition; John Mayow and Robert Boyle 170
# The Aristotelian Balance—sheet
# Aristotle’s Theory of Causation
# The Hellenistic Age
# Galen and the Vital Faculties
[[Book - A History of Embryology 1959-2|'''Chapter Two - Embryology From Galen To The Renaissance''']]
# Patristic Speculation
# Contributions of Jewish Thinkers
# Embryology among the Arabs
# Alchemy and Embryology
# The Visions of St Hildegard
# Albertus Magnus; the Re-awakening of Scientific Embryology
# Aristotle’s Masterpiece
# Scholastic Ideas on Generation
# The Insights of Leonardo da Vinci
# The Macro-Iconogrnphers of the Sixteenth Century
# The Movement to Rationalise Obstetrim
[[Book - A History of Embryology 1959-3|'''Chapter Three - Embryology In The Seventeenth Century''']]
# The Opening Years
# Developmental Determinism and Trasplantation; Digby, Higlunom and Tagliacozzi
# Thomas Browne and the Beginnings of Chemical Embryology
# William Harvey and the Identification of the Blastoderm
# The Riddle of Fertilisation
# Harvey's Achievements and Influence
# Atomist Theories of Embryonic Development; Gaucndi and Dcscm-ta
# Fixatives and Uterine Milk; Robert Boyle and Walter Needhzun
# The Discovery of the Follicles of the Mammalian Ovary
# The Micro-iconographers and Preformatianism; Marcello Malpighi and Jan Swammerdam
# Foetal Respiration and Composition; John Mayow and Robert Boyle
[[Book - A History of Embryology 1959-4|'''Chapter Four - Embryology In The Eighteenth Century''']]
# Theories of Foetal Nutrition
# Growth and Difierentiation; Stahl and Main-e-Jan
# Chemical and Quantitative Approaches to the Origin of Organisation; Boerhnave, Hamberger and Mazin
# Albrecht van Haller and the Rise of Techniques
# Embryos and Theologians
# Ovism and Animzdculism
# Spontaneous Generation
# Preformation and Epigenesis
# The Closing Years
[[Book_-_A_History_of_Embryology_1959-4#Conclusion|'''Conclusion''']]


Chapter Four
[[Book_-_A_History_of_Embryology_1959-4#Bibliography|'''Bibliography''']]
EMBRYOLOGY IN THE EIGHTEENTH CENTURY


1. Theories of Foetal Nutrition 179
2. Growth and Difierentiation; Stahl and Main-e-Jan 183
3. Chemital and Quantitative Approaches to the Origin of Organisation; Boerhnave, I-lunberger and Mazin I86
4. Albrecht van Haller and the Rise of Techniques :93
5. Embryos and Theologians 204
6. Ovism and Animzdculism 205
7. Spontaneous Generation 2!!
8. Prcforrmrion and Epigenesis 213
9. The Closing Years 223


Conclusion 230
Traditional methods of incubation
Bibliography 241
Index 293
I Traditional methods of incubation


Portrait of William Harvey, act. 6t (1639)
Portrait of William Harvey, act. 6t (1639)


PLATES
==Preliminary Note==


(A) Egyptian peasant incubator (from Cadman)
The contribution to the history of science contained in the following four chapters first appeared as the opening part of a treatise on Chemical Embryology, published in 1931. They were delivered in the form of lectures about the same time at the University of London under the title “Speculation, Observation and Experiment as illustrated by the History of Embryology." The munificence of that University assured their appearance in separate, and amplified, form.‘
(B) Chinese peasant incubator (from King)


Heteromorphosis in a guardian deity (lokapfila) depicted
I suppose that the study of the history of science needs no apology. If at first sight the discussion of what was thought in the past rather than what is known now appmrs to be of merely antiquarian value, a deeper consideration will admit, with Louis Choulant, that the history of science is the guarantee of its freedom. The mistakes of our predecessors remind us that we may be mistaken; their wisdom prevents us from assuming that wisdom was born withus; and by studying the processes of their thought, we may hope to have a better understanding, and hence a better organisation, of our own. Theoreticil errors, such as the final cause, preformationisrn or phlog-iston; practical errors, such as the divorce between speculation and technique in the Hellenistic age, are always able to show us a more excellent way.
in a fresco on the wall of one of the m‘vc~tempIes at
Ch’ien-F0-Tung, Tunlnmng, Kansu province, China


The oldest known drawing of the uterus. From a ninth-
The present contribution does not claim, what probably no historical work can tnrly deserve, the ascription of a complete lack of bias in its presentation. Designed as it was to introduce a discussion of the border- line between embryology and biochemistry, it sought rather to lay bare the roots of chemical embryology in history, than to collect data indis- criminately on all the interesting aspects of the subject. Its title, “The Origins of Chemical Embryology," made no secret of this. And no obvious disadvantage attaches to such a plan, except the difficulty of deciding when to leave off. For although it is possible in reasonable space to try to dojustice to all aspects of embryology before 1800, after that date the number of investigators and thevariety of problems attacked becomes too great to handle conveniently on the same scale as before.’
century MS. (the Brussels Mosdlion Codex) of Soranus’
work on gynaecology


An illustration from the Liber Sn'1:z'as (LAD. 1150) of St
‘ By embryology we mean in this book the embryology of animal: txduxivlly. The history ofthe embryology of plant: has been fully written only in Russian, by Bannov, but there is 1 shorter work by Souéges in French.
Hildegard of Bingen (Wicsbaden Codex B), showing the
descent of the soul into the embryo (after Singer)


A page from Leonardo da Vinci’s anatomical notebooks
I or. the valuable work of Srudmflu; Florian; Dogelb; Oppenheimer; Fischer 5: Schopfer; and others.
(guaderni d’Anazmm'a), c. AD. 1490


Portrait of Volcher Coiter, act. 41 (painted in r 575 byan
Bifurcation bcgms; the spheres of morphology and ph,11o1ogy more obviously separate, and In the latter division chemical researches play an ever-increasing part. It is now hoped that a group of workers Wlll soon be able to continue the story in a companion volume through the nineteenth century under a number of separate hadings.
unknown master)


Portrait of Sir Kenelm Digby (from the painting by
No exhaustive treatise on the history of embryology as yet exists.‘
Oomelius Jansen, ca. 1650)
 
Portrait of Sir Thomas Browne and his wife,
Dorothy (ca. x650)
 
IX Zeus liberating living beings from an egg (the frontispieoe
 
of William Harvey’s book on the Generation of Animal:,
1 6 51)
 
X Illustrations from “falter Needl1am’s De Fonnatn Foetu of 1667
 
Fmntirpieze
 
facing page 24
 
84
 
96
 
m4
 
122
 
132
 
‘33
 
:58
XII
 
XIII
 
XVII
 
XVII!
 
PLATES
 
Iflustrations from Maipighfs 1): 01:0 Incubate of 167:
shgwmg the early stages of the development of the
°h“'-k facing page 166
 
Illustrations from 77:: auiou: and accurate ab.mvan'an: 0/
Mr Stephen Lorene-t'm' of Florence on the Dirxrrtioru qf
 
the Cramp-Fish ,9,
Metamorphosis in Buddhist iconography; statues in the
 
Sleeping Buddha Temple at Suchaw (Chin-dffian),
 
Kansu province. China ,7;
Portrait at‘ Antan van Leeuwmhoek by Job.


Vex-kalje (1686) :7:
The nearest approach to it is the very valuable memoir of E. Bloch with
Portrait of Robert Boyle (a. 1690) 176
its epitome. but this only covers the era of the Renaissance with thor- oughness. Hertwig's account, which he printed at the beginning of his great Haruibuth tier Enlwicfalungxlehre, does not deal very fully with any aspect of the subject before x8oo, nor do the much shorter ones of Hen- neguy and Minot. The latter paper is interesting in that it ends with an emphasis on the need for physico-cherniczil work in the future. The introduction to Keibel's book is much slight;-r, but contains some useful information. There are various monographs and papers on special points, such as Youchet's rather untrustworthy treatment of the em- bryology of Aristotle, and Lones’ discussion of it, which is worse. Camus‘ notes are still the best commentary on the I1z'.rtlm‘a Animalium. Again, useful information on some cultural points is to be had from the treatise ofl"loss 6: Bartels. The introductions to certain books also con- tain valuable information, and in this class comes Dareste's remarkable book on temtology. The bibliographies contained in Von Hallefs eighth volume and in the books of Schurig and Hefiter are naturally of the greatest assistance. The valuable books of F. J. Cole and Thadeusz Bilikiewicz on seventeenth~century embryology appeared too late for use in the first prepamtion of this book, but have contributed to its revision.’
A tel-atoms with well-formed teeth and hair (from Robert


Plat‘: Natural Hi:-tmy afS!ufl'ord:h:'r:, 1686) 178
In 1939 there appeared a work, The Rise of Ermlryalagy, by the learned Californian anatomist A. W. Meyer, author of numerous periodical publimtions on our subject, some of which are referred to in the bibliography. His book stands to mine in much the same relation as the second volume of David Eugene Smith’s notable History qfMaihe- malice to the first; the one adopting 2| basically chronological treatment, the other A topical form in which separate subjects are chosen in succes- sion for consideration. However, Meyer devotes the bulk of his work to
Illustrations from Antoine Maine-Jan‘: Obaematiovu


mr Infmnahbn du pcmlet (1722) 186
I am we cannot attempt to provide a bibliography of the more upP°¢,m'fl modem work dealing with the subject melt". Yet in case suenufic mm or historians of other fields might appreciate some helpful introduction to embryology. Inlntton rnly ‘>9 made oi the popular boolu of Rouund, Wnddington mil Guttnnrher. An engineer or on hillflnan of xstronmny might lhlfl proceed to the recent tunes: of Wlddington,
De Réaumux-‘I Incubators (from De (‘art de faire (chm
 
In paulm, 1749) 204


N O T E The use of the Ihonened and (&) indium eollzbcmion between two or more author
man, or Willier 2: al. _ . - Certain minor works on the history of embryology hive proved maee¢sxble— Beulre; Eccleahymer; H. Fnheuder; Fnvaro; l’-‘enzkel; Gills: Hapr; Ottow. Other
 
PRELIMINARY NOTE
 
THE contribution to the history of science contained in the following
four chapters first appeared as the opening part of a treatise on Chemical
Embryology, published in 1931. They were delivered in the form of
lectures about the same time at the University of London under
the title “Speculation, Observation and Experiment as illustrated by
the History of Embryology." The munificence of that University
assured their appearance in separate, and amplified, form.‘
 
I suppose that the study of the history of science needs no apology.
If at first sight the discussion of what was thought in the past rather than
what is known now appmrs to be of merely antiquarian value, a deeper
consideration will admit, with Louis Choulant, that the history of
science is the guarantee of its freedom. The mistakes of our predecessors
remind us that we may be mistaken; their wisdom prevents us from
assuming that wisdom was born withus; and by studying the processes
of their thought, we may hope to have a better understanding, and hence
a better organisation, of our own. Theoreticil errors, such as the final
cause, preformationisrn or phlog-iston; practical errors, such as the
divorce between speculation and technique in the Hellenistic age, are
always able to show us a more excellent way.
 
The present contribution does not claim, what probably no historical
work can tnrly deserve, the ascription of a complete lack of bias in its
presentation. Designed as it was to introduce a discussion of the border-
line between embryology and biochemistry, it sought rather to lay bare
the roots of chemical embryology in history, than to collect data indis-
criminately on all the interesting aspects of the subject. Its title, “The
Origins of Chemical Embryology," made no secret of this. And no
obvious disadvantage attaches to such a plan, except the difficulty of
deciding when to leave off. For although it is possible in reasonable
space to try to dojustice to all aspects of embryology before 1800, after
that date the number of investigators and thevariety of problems attacked
becomes too great to handle conveniently on the same scale as before.’
 
‘ By embryology we mean in this book the embryology of animal: txduxivlly. The
history ofthe embryology of plant: has been fully written only in Russian, by Bannov,
but there is 1 shorter work by Souéges in French.
 
I or. the valuable work of Srudmflu; Florian; Dogelb; Oppenheimer; Fischer 5:
Schopfer; and others.
 
Bifurcation bcgms; the spheres of morphology and ph,11o1ogy
more obviously separate, and In the latter division chemical researches
play an ever-increasing part. It is now hoped that a group of workers
Wlll soon be able to continue the story in a companion volume through
the nineteenth century under a number of separate hadings.
 
No exhaustive treatise on the history of embryology as yet exists.‘
:I'he nearest approach to it is the very valuable memoir of E. Bloch with
its epitome. but this only covers the era of the Renaissance with thor-
oughness. Hertwig's account, which he printed at the beginning of his
great Haruibuth tier Enlwicfalungxlehre, does not deal very fully with any
aspect of the subject before x8oo, nor do the much shorter ones of Hen-
neguy and Minot. The latter paper is interesting in that it ends with an
emphasis on the need for physico-cherniczil work in the future. The
introduction to Keibel's book is much slight;-r, but contains some useful
information. There are various monographs and papers on special
points, such as Youchet's rather untrustworthy treatment of the em-
bryology of Aristotle, and Lones’ discussion of it, which is worse.
Camus‘ notes are still the best commentary on the I1z'.rtlm‘a Animalium.
Again, useful information on some cultural points is to be had from the
treatise ofl"loss 6: Bartels. The introductions to certain books also con-
tain valuable information, and in this class comes Dareste's remarkable
book on temtology. The bibliographies contained in Von Hallefs eighth
volume and in the books of Schurig and Hefiter are naturally of the
greatest assistance. The valuable books of F. J. Cole and Thadeusz
Bilikiewicz on seventeenth~century embryology appeared too late for
use in the first prepamtion of this book, but have contributed to its
revision.’
 
In 1939 there appeared a work, The Rise of Ermlryalagy, by the
learned Californian anatomist A. W. Meyer, author of numerous
periodical publimtions on our subject, some of which are referred to in
the bibliography. His book stands to mine in much the same relation as
the second volume of David Eugene Smith’s notable History qfMaihe-
malice to the first; the one adopting 2| basically chronological treatment,
the other A topical form in which separate subjects are chosen in succes-
sion for consideration. However, Meyer devotes the bulk of his work to
 
I am we cannot Ittempt to pmvide . bibliography of the more upP°¢,m'fl modem
worh dealing with the subject melt". Yet in case suenufic mm or historians of other
fields might appreciate some helpful introduction to embryology. Inlntton rnly ‘>9
made oi the popular boolu of Rouund, Wnddington mil Guttnnrher. An engineer or
on hillflnan of xstronmny might lhlfl proceed to the recent tunes: of Wlddington,
 
man, or Willier 2: al. _ .
- Certain minor works on the history of embryology hive proved maee¢sxble—
Beulre; Eccleahymer; H. Fnheuder; Fnvaro; l’-‘enzkel; Gills: Hapr; Ottow. Other


urtides deserving mention are those of Gerber; Keller; du Ems.
urtides deserving mention are those of Gerber; Keller; du Ems.




the eighteenth and nineteenth centuries, passing over the earlier periods
the eighteenth and nineteenth centuries, passing over the earlier periods in his first thirty pages. His treatment of the nineteenth century is in- teresting indeed, though nothing could supersede the remarkable work of E. S. Russell, Form and Function. Particular interest attaches to L. A. Blacher’s monograph on Embryology in Russia in the 18!]: and 191/: Centun'e.r (195 5), since so much of the classical work centring around 1800 was done or published in that country.
in his first thirty pages. His treatment of the nineteenth century is in-
teresting indeed, though nothing could supersede the remarkable work
of E. S. Russell, Form and Function. Particular interest attaches to L. A.
Blacher’s monograph on Embryology in Russia in the 18!]: and 191/:
Centun'e.r (195 5), since so much of the classical work centring around
1800 was done or published in that country.


These observations made, the principal reviews of the subject are
These observations made, the principal reviews of the subject are chiefly to be found in histories of science in general, such as Sartou’s; histories of biological theory, such as R.’idl’s; histories of obstetrics, such as von SieboId's, Spencer's and E. F asbender's; histories ofgynaecology, such as McKay’s; and histories of anatomy, such as Singer's and Von T6ply's. Histories of medicine as a whole are numerous and good: I have found those of Garrison and Neuburgcr-Pagel most useful. Those which deal with special periods are also of assistance, such as Schrutz and Browne on Arabian, I. Bloch on Byzantine, and I-larnack on Patristic medicine. Histories of chemistry provide no help, for ancient chemistry was so oriented towards “practical” results, such as the lapi: philorophomm and elixir vitae, that the egg was only considered as a raw material for various preparations. The investigation of its change of properties during the development of the embryo did not occur to the alchemists. Detailed studies of particular subjects, such as those con- tained in Singer's two excellent volumes, The History and Method of Science, may also be of some assistance. Again, there are books which give a wonderful orientation and an articulate survey of vast tracts: of these Clifford Allbutt’s Greek Medicine in Rome, with its mass of refer- ences, is among the most valuable. And Miall’s Earbv Naturalist: must not be omitted, far, apart from the peculiar charm of style which marks it, it contains some singularly helpful bibliographical data.‘ But the study of the original sources, so far as that is possible, is a duty which cannot be avoided, and in what follows I have been careful to copy down no statement from a previous review when it was possible to read the actual words of the writer himself. This practice of going to the originals is made peculiarly necessary in a case such as the present one, when the history of a subject is regarded from a rather new angle.
chiefly to be found in histories of science in general, such as Sartou’s;
histories of biological theory, such as R.’idl’s; histories of obstetrics, such
as von SieboId's, Spencer's and E. F asbender's; histories ofgynaecology,
such as McKay’s; and histories of anatomy, such as Singer's and Von
T6ply's. Histories of medicine as a whole are numerous and good: I
have found those of Garrison and Neuburgcr-Pagel most useful. Those
which deal with special periods are also of assistance, such as Schrutz
and Browne on Arabian, I. Bloch on Byzantine, and I-larnack on
Patristic medicine. Histories of chemistry provide no help, for ancient
chemistry was so oriented towards “practical” results, such as the lapi:
philorophomm and elixir vitae, that the egg was only considered as a raw
material for various preparations. The investigation of its change of
properties during the development of the embryo did not occur to the
alchemists. Detailed studies of particular subjects, such as those con-
tained in Singer's two excellent volumes, The History and Method of
Science, may also be of some assistance. Again, there are books which
give a wonderful orientation and an articulate survey of vast tracts: of
these Clifford Allbutt’s Greek Medicine in Rome, with its mass of refer-
ences, is among the most valuable. And Miall’s Earbv Naturalist: must
not be omitted, far, apart from the peculiar charm of style which marks
it, it contains some singularly helpful bibliographical data.‘ But the
study of the original sources, so far as that is possible, is a duty which
cannot be avoided, and in what follows I have been careful to copy down
no statement from a previous review when it was possible to read the
actual words of the writer himself. This practice of going to the
originals is made peculiarly necessary in a case such as the present one,
when the history of a subject is regarded from a rather new angle.


The arrangement of my chapters I adopted in the first edition, and
The arrangement of my chapters I adopted in the first edition, and now preserve, only on the ground that it is suitable enough in the pres~ ent state of historical knowledge. Little was then said about eml>ry~ ology in China beuxusc at that time I could find out little about it, but it will be thoroughly treated in the eighth volume of my work on the
now preserve, only on the ground that it is suitable enough in the pres~
ent state of historical knowledge. Little was then said about eml>ry~
ology in China beuxusc at that time I could find out little about it, but
it will be thoroughly treated in the eighth volume of my work on the


‘ A fine beginning has been made on the bibliographies of seventeenth-century mm
‘ A fine beginning has been made on the bibliographies of seventeenth-century mm of science by Keyna and Fulton.
of science by Keyna and Fulton.


history-of science in general in that great culture, Scimte and Ciril-
history-of science in general in that great culture, Scimte and Ciril- "“"""_ W C{'1"0- Nor am I content with the short section on embry- ology In India, but here there are special difficulties owing to thc absmce of an established chronology for ancient and mediaeval Indian texts and an adequate account of it must be left for others to give. No per: manent framework for historical facts is proposed in what follows; I only attempt to bring them together, and to reveal some of the relation. ships between them. If the traditional pattern turns out to be badly distorted-—and there are many signs that it may—the facts can be rearranged.
"“"""_ W C{'1"0- Nor am I content with the short section on embry-
ology In India, but here there are special difficulties owing to thc absmce
of an established chronology for ancient and mediaeval Indian texts
and an adequate account of it must be left for others to give. No per:
manent framework for historical facts is proposed in what follows; I
only attempt to bring them together, and to reveal some of the relation.
ships between them. If the traditional pattern turns out to be badly
distorted-—and there are many signs that it may—the facts can be
rearranged.


But in whatever way this may tum out to be desirable, one neeasity
But in whatever way this may tum out to be desirable, one neeasity rnust constantly be kept before the mind's eye, namely the knowledge of the relations between scientific thought and technical practice at any given period. For embryology this knowledge is diflicult to acquire, since up to the time of the Renaissance obstetrics remained a part of primitive folk-medicine rather than of serious medical science. We see, however, in the publication of the Hellenistic gynaecological treatises in the sixteenth century (Bauhin, Spach; see p. 109) the satisfaction of a new demand, even though it took the typiml Renaissance form of what might be mlled palaeolatry. It was part of that movement to rationalise obstetric: which included Harvey’s De Gmnatione and Malpighi's De Formatione Pulli and culminated in the celebrated man-midwives of the eighteenth century.‘ Again, the relation of the early systematists— Belon, Rondelet, Aldrovandus, Ray-—to the beginnings of mermntile expansion is fairly clear, for the mediaeval bstiary could not cope with the influx of new animals and plants from hitherto unknown regions, any one of which might prove to be an exploitable commodity.
rnust constantly be kept before the mind's eye, namely the knowledge of
the relations between scientific thought and technical practice at any
given period. For embryology this knowledge is diflicult to acquire,
since up to the time of the Renaissance obstetrics remained a part of
primitive folk-medicine rather than of serious medical science. We see,
however, in the publication of the Hellenistic gynaecological treatises in
the sixteenth century (Bauhin, Spach; see p. 109) the satisfaction of a
new demand, even though it took the typiml Renaissance form of what
might be mlled palaeolatry. It was part of that movement to rationalise
obstetric: which included Harvey’s De Gmnatione and Malpighi's De
Formatione Pulli and culminated in the celebrated man-midwives of the
eighteenth century.‘ Again, the relation of the early systematists—
Belon, Rondelet, Aldrovandus, Ray-—to the beginnings of mermntile
expansion is fairly clear, for the mediaeval bstiary could not cope with
the influx of new animals and plants from hitherto unknown regions,
any one of which might prove to be an exploitable commodity.


The Hellenistic divorce between scientific thought and empirical
The Hellenistic divorce between scientific thought and empirical technique is an important case in point. Greek life was divided strictly ima amela and ngdftg. The latter was not thought fitting [or a man of good birth. "Antiquity," says Diels. “was entirely aristocratic in attitude. Even prominent artists. Such 35 P5555335. W619 0135355 33 fifti- sans, and were incapable of bursting through the barrier separating the workers and pmsants from the upper clam. A second muse of the slight technical progress in antiquitywas its slave-holding system, which led to a lack of any impulse to develop the machine as a substitute for manual labour.” Xenophon in the Oemnomicus held the industries in poor repute.‘ "Men engaged in the mechanical arts,” he says, “must ever be
technique is an important case in point. Greek life was divided strictly
ima amela and ngdftg. The latter was not thought fitting [or a man
of good birth. "Antiquity," says Diels. “was entirely aristocratic in
attitude. Even prominent artists. Such 35 P5555335. W619 0135355 33 fifti-
sans, and were incapable of bursting through the barrier separating the
workers and pmsants from the upper clam. A second muse of the slight
technical progress in antiquitywas its slave-holding system, which led to
a lack of any impulse to develop the machine as a substitute for manual
labour.” Xenophon in the Oemnomicus held the industries in poor
repute.‘ "Men engaged in the mechanical arts,” he says, “must ever be


‘!3.g. the Chnmberlenx. Palfyn (see Stein), Mluricuu, _VVfl.ltarn Srnellrz, John
‘!3.g. the Chnmberlenx. Palfyn (see Stein), Mluricuu, _VVfl.ltarn Srnellrz, John Burton of York ("Dr Slop"), and Joseph Needhnm or Dewnzes; see the Imeles or Rosemhnl and Mengert. The dissertation ofCaspar Base (1729) is I typuml attack on the midwives ufllll time.
Burton of York ("Dr Slop"), and Joseph Needhnm or Dewnzes; see the Imeles or
Rosemhnl and Mengert. The dissertation ofCaspar Base (1729) is I typuml attack on
the midwives ufllll time.


‘ See Creeotti. ' xv. 3; VI. lJ'l5-
‘ See Creeotti. ' xv. 3; VI. lJ'l5-


both bad friends and feeble defenders of their country." He troubled
both bad friends and feeble defenders of their country." He troubled himself little with those skilful in carpentry, metallurgy, painting and sculpture, but was always anxious to meet. a “gentleman" (ualzig rs miyafldg). The results of this were inevitable. Classical surgery and obstetrics benefited practically nothing from the speculations of the biologists from Alcmaeon to Herophilus (see pp. 29 Surgeons and midwives remained members of the painter-cobbler-builder group, the group of base-bom "rnechanicks”, entirely distinct from the astronomer- mathematician-xnetaphysician-biologist group, the group familiar with courts and tyrants.
himself little with those skilful in carpentry, metallurgy, painting and
sculpture, but was always anxious to meet. a “gentleman" (ualzig rs
miyafldg). The results of this were inevitable. Classical surgery and
obstetrics benefited practically nothing from the speculations of the
biologists from Alcmaeon to Herophilus (see pp. 29 Surgeons and
midwives remained members of the painter-cobbler-builder group, the
group of base-bom "rnechanicks”, entirely distinct from the astronomer-
mathematician-xnetaphysician-biologist group, the group familiar with
courts and tyrants.
 
Only the greatest broke away from this tradition: Aristotle, when he
conversed with fishermen, Archimedes perhaps, when he constnicted
his mechanical devices. For the rest, it was too strong. Down to the end
of the Roman period the artillery in use remained precisely what it had
been six hundred years before, although the Empire was crumbling
under barbarian pressure, and would have given anything, one would
imagine, for an improved artillery capable of withstanding the Gothic
armies. It is strange, as has been acutely said, that the Rornans never
invented anything so much in the Roman taste as a railway. So far as
Hellenistic empirical industrial chemistry was concerned, the Demo-
critcan and Epicurean atoms might never have existed. And in medicine,
the only effect of the brilliant Greek atomic speculations was to give rise
to the Methodic school of Roman physicians, described by Allbutt,
whose influence was never strong, and who contributed relatively little
to the main stream of therapeutics originating with Hippocrates.


In sum, we must not dissociate scientific advances from the technical
Only the greatest broke away from this tradition: Aristotle, when he conversed with fishermen, Archimedes perhaps, when he constnicted his mechanical devices. For the rest, it was too strong. Down to the end of the Roman period the artillery in use remained precisely what it had been six hundred years before, although the Empire was crumbling under barbarian pressure, and would have given anything, one would imagine, for an improved artillery capable of withstanding the Gothic armies. It is strange, as has been acutely said, that the Rornans never invented anything so much in the Roman taste as a railway. So far as Hellenistic empirical industrial chemistry was concerned, the Demo- critcan and Epicurean atoms might never have existed. And in medicine, the only effect of the brilliant Greek atomic speculations was to give rise to the Methodic school of Roman physicians, described by Allbutt, whose influence was never strong, and who contributed relatively little to the main stream of therapeutics originating with Hippocrates.
needs and processes of the time, and the economic structure in which all
are embedded. We shall never understand the failure of Greek science
if we consider it in abstraction from the environment which sterilised its
speculation. The history of science is not a mere succession of in-
explicable geniuses, direct Promethean ambassadors to man from
heaven. Vlfhether a given fact would have got itself discovered by some
other person than the historical discoverer had he not lived, it is cer-


tainly profitless and probably meaningless to enquire. But scientific
In sum, we must not dissociate scientific advances from the technical needs and processes of the time, and the economic structure in which all are embedded. We shall never understand the failure of Greek science if we consider it in abstraction from the environment which sterilised its speculation. The history of science is not a mere succession of in- explicable geniuses, direct Promethean ambassadors to man from heaven. Whether a given fact would have got itself discovered by some other person than the historical discoverer had he not lived, it is certainly profitless and probably meaningless to enquire. But scientific men do not live in a vacuum; on the contrary, the directions of their interest are ever conditioned by the structure of the world they live in. Further historical research will enable us to take into account the social and economic status of the invatigator himself (cf. Chambers for the Hellenistic artist, and Yearsley for the sixteenth-century physician).
men do not live in a vacuum; on the contrary, the directions of their
interest are ever conditioned by the structure of the world they live in.
Further historical research will enable us to take into account the social
and economic status of the invatigator himself (cf. Chambers for the
Hellenistic artist, and Yearsley for the sixteenth-century physician).


It would thus be of the greatest interest to lmow accurately the sources
It would thus be of the greatest interest to lmow accurately the sources of the emoluments of embryologists at different times.‘ From Om-
of the emoluments of embryologists at different times.‘ From Om-


‘ On this, cf. Cumston and Dittrielr.
‘ On this, cf. Cumston and Dittrielr.


steinjstadmirzible hook on the scientific societies of the Renaissance, the
steinjstadmirzible book on the scientific societies of the Renaissance, the suspicion arises that their royal patronage was dictated not only by
suspicion arises that their royal patronage was dictated not only by
:1 purely disinterested passion {or abstract truth, but by a desire to profit
as much as possible by the new techniques which the decay of the gnfi-
usury doctrines, the Willingness of the rising mercantile class m mkc
mdusmfil "ffimlllts." and um f”'1'3“8i"8 thought of the scientific men
were combining to produce. In England's Royal Society, indeed, me
preoccupation of the early Fellows with the Uirnprovexnent of u-ad; and
husbandry” is patent to anyone acquainted with its early history (cf.
Thomas Spint’s account of it)! Thus Dr Jasper Needham, clccled in
1663, read only one paper before the Society—not, as might haw; been
expected from his profession, on the transfusion of blood or the anatomy
of the brain; but on the value and use of “China Varnish". However, it
is probable that for the most part the emhryologists whose work we
shall have to discuss were pi-.iciisi.ng physicians, free or relatively free
from the ancient tnidition, and conscious that to understand the mystery
of generation would be to advance the science and art of medicine.


In this connection it is of interest that the Church in the seventeenth
1 purely disinterested passion {or abstract truth, but by a desire to profit as much as possible by the new techniques which the decay of the gnfi- usury doctrines, the Willingness of the rising mercantile class m mkc mdusmfil "ffimlllts." and um f”'1'3“8i"8 thought of the scientific men were combining to produce. In England's Royal Society, indeed, me preoccupation of the early Fellows with the Uirnprovexnent of u-ad; and husbandry” is patent to anyone acquainted with its early history (cf. Thomas Spint’s account of it)! Thus Dr Jasper Needham, clccled in 1663, read only one paper before the Society—not, as might haw; been expected from his profession, on the transfusion of blood or the anatomy of the brain; but on the value and use of “China Varnish". However, it is probable that for the most part the emhryologists whose work we shall have to discuss were pi-.iciisi.ng physicians, free or relatively free from the ancient tnidition, and conscious that to understand the mystery of generation would be to advance the science and art of medicine.
and eighteenth centuries provided a certain source of demand for em-
bryological research. Of this Swaminerdarn and Malehranche (see
p. x69) provide interesting examples, and the conviction, then widely
held, that research into the nature of generation would throw light an
orthodox theological doctrines, such as that of original sin, led to an
economic situation of value for biological development. Finally, it
would he rash to minimise the factor of pure curiosity in seventeenth-
century science. The recreational quality of Lecuwenhoek's investiga-
tions is, as Baas-Becking says, too obvious to be overlooked.


The history of single forms of scientific knowledge is in way hap-
In this connection it is of interest that the Church in the seventeenth and eighteenth centuries provided a certain source of demand for em- bryological research. Of this Swaminerdarn and Malehranche (see p. x69) provide interesting examples, and the conviction, then widely held, that research into the nature of generation would throw light an orthodox theological doctrines, such as that of original sin, led to an economic situation of value for biological development. Finally, it would he rash to minimise the factor of pure curiosity in seventeenth- century science. The recreational quality of Lecuwenhoek's investiga- tions is, as Baas-Becking says, too obvious to be overlooked.
pier because containing more of continuity than that of civilisation as a
whole. The assiduity with which men of diflerent periods in the rise and
decline of a culture pursue the diflerent forms of human experience may,
as Spengler has shown, vary much, but those forms remain funda-
mentally the same, even if their manifestations are profoundly changed.


The history of single forms of scientific knowledge is in way hap- pier because containing more of continuity than that of civilisation as a whole. The assiduity with which men of diflerent periods in the rise and decline of a culture pursue the diflerent forms of human experience may, as Spengler has shown, vary much, but those forms remain funda- mentally the same, even if their manifestations are profoundly changed.


scphy, the mechanic, and husbandry, according to the principles of our new philoso-
phical college, that Values no knowledge, but as it hath a tendency in use.And therefore
I shall make K on: of my xuiu to you, that you would take the pains to enquire lllILl€
more thoroughly inro the way: of husbandry etc. practised in your pan}: ‘rid when you
intend for England, to bring along with you what good receipt: or choice books of my
of those subjects you can procure; which will make you extremely welcome to our


‘ ‘b I , hichlhad d ' edt ‘ oulil 'ti of.”Fultonre-
scphy, the mechanic, and husbandry, according to the principles of our new philoso- phical college, that Values no knowledge, but as it hath a tendency in use.And therefore I shall make K on: of my xuiu to you, that you would take the pains to enquire lllILl€ more thoroughly inro the way: of husbandry etc. practised in your pan}: ‘rid when you intend for England, to bring along with you what good receipt: or choice books of my of those subjects you can procure; which will make you extremely welcome to our
innilxlhlihillttiflh imzmmi of nuieiiifé but wisgiapy taaliie leave to think


it was not to inadequate as um-rywould suppose. _ .
‘ ‘b I , hichlhad d ' edt ‘ oulil 'ti of.”Fultonre- imzmmi of nuieiiifé but wisgiapy taaliie leave to think
- The run loop: of Leeuwenhoek‘: discoveries Is now appearing, drum to the


labour: of van Runberlc and his mllabontvfi.
it was not to inadequate as um-rywould suppose. _ . - The run loop: of Leeuwenhoek‘: discoveries Is now appearing, drum to the labour: of van Runberlc and his mllabontvfi.




That science, at any rate, does maintain some sort of continuity what-
That science, at any rate, does maintain some sort of continuity what- ever gaps there may be between the phases of its progress, is a belief agreeable with all the available facts, and one which no criticism will easily shake.
ever gaps there may be between the phases of its progress, is a belief
agreeable with all the available facts, and one which no criticism will
easily shake.


It only remains to record my indebtedness to those who have assisted
It only remains to record my indebtedness to those who have assisted me in the preparation of this work. Primarily I am grateful to Dr Charles Singer, who annotated my typescript with valuable comments and lent me many papers and pictures, and to Professor R. C. Punnett, who placed unreservedly at my disposal his knowledge of the history of generation and his library of old and rare biological books. To Dr Arthur Peck I am indebted for the correction of my Greek, and it was Professor A. B. Cook who introduced me to the embryology of the ancients. For guidance on Talmudic and Jewish matters I thank Dr Walter Pagel, the late Dr Louis Rapkine and Dr H. Loewe. Without the assiduous backing of Mr Powell, the Librarian of the Royal Society of Medicine, and his assistants, and of Mr H. Zeitlinger, I should have dealt much more inadequately than I have with papers and books which cannot be consulted in Cambridge. And in addition to those mentioned above, the following friends kindly read through and criticised the proofs: Pro- fessor Reuben Levy, the late Professor F. M. Cornford, the late Sir William Dampier, Mr Gregory Bateson, Professor Roy Pascal and the Rev. W. L. Elmslie.
me in the preparation of this work. Primarily I am grateful to Dr Charles
Singer, who annotated my typescript with valuable comments and lent
me many papers and pictures, and to Professor R. C. Punnett, who
placed unreservedly at my disposal his knowledge of the history of
generation and his library of old and rare biological books. To Dr Arthur
Peck I am indebted for the correction of my Greek, and it was Professor
A. B. Cook who introduced me to the embryology of the ancients. For
guidance on Talmudic and Jewish matters I thank Dr Walter Pagel,
the late Dr Louis Rapkine and Dr H. Loewe. Without the assiduous
backing of Mr Powell, the Librarian of the Royal Society of Medicine,
and his assistants, and of Mr H. Zeitlinger, I should have dealt much
more inadequately than I have with papers and books which cannot be
consulted in Cambridge. And in addition to those mentioned above, the
following friends kindly read through and criticised the proofs: Pro-
fessor Reuben Levy, the late Professor F. M. Cornford, the late Sir
William Dampier, Mr Gregory Bateson, Professor Roy Pascal and the
Rev. W. L. Elmslie. _


To the Master of Gonville and Caius College I am indebted for
permission to reproduce the portrait of William Harvey (attributed to
Rembrandt) which hangs in our Senior Combination Room. Although
the authenticity of this is not accepted by Keynes in his recent study of
the portraits of Harvey, it has been in the possession of the College
since r798, when it came to us from the Earl of Leicester. After com-
parison with other portraits of Harvey, many feel unable to concur in its
rejection.


==Chapter 4==
To the Master of Gonville and Caius College I am indebted for permission to reproduce the portrait of William Harvey (attributed to Rembrandt) which hangs in our Senior Combination Room. Although the authenticity of this is not accepted by Keynes in his recent study of the portraits of Harvey, it has been in the possession of the College since r798, when it came to us from the Earl of Leicester. After com- parison with other portraits of Harvey, many feel unable to concur in its rejection.
EMBRYOLOGY IN THE EIGHTEENTH CENTURY




r. Theories of Foetal Nutrition -
==Plates==


DURING the course of the seventeenth, and the _fi_rs_t quarter of the
(A) Egyptian peasant incubator (from Cadman)
eighteenth, century, many theories were propounded concei-ning foetal
(B) Chinese peasant incubator (from King)
nutrition. It is convenient to classify them (Table I).


At this point the Emmenalogia of John Freiud' deserves special
Heteromorphosis in a guardian deity (lokapfila) depicted
reference. This was a book which dealt with all aspects of menstruation.’
in a fresco on the wall of one of the m‘vc~tempIes at
As has already been mentioned (p. 15o n. 2) be supposed that the blood
Ch’ien-F0-Tung, Tunlnmng, Kansu province, China
passing through the placenta to the embryo was distinctively menstrual
blood. This view he supported by an arithmetical argument. Calculating
the amount of menstrual blood evacuated in nine months, he said,


The quantity of Blood which the Mother may bestow upon the nourish-
The oldest known drawing of the uterus. From a ninth-
mentof her Ofispringwillbe lib. 13 nzt.2§,whichwill outweigh the newborn
century MS. (the Brussels Mosdlion Codex) of Soranus’
Foetus with all its Integurnents, if they should be put into a balance; and
work on gynaecology


' leave no room to doubt, its being able to bestow very proper nourishment on
An illustration from the Liber Sn'1:z'as (LAD. 1150) of St
Hildegard of Bingen (Wicsbaden Codex B), showing the
descent of the soul into the embryo (after Singer)


the Embrio. For the mean weight of a new-bom Foetus is about 12 lx'b.,
A page from Leonardo da Vinci’s anatomical notebooks
sometirns it is found greater, and very often less.
(guaderni d’Anazmm'a), c. AD. 1490


This quantitative outlook forms a parallel to Harvey's approach in his
Portrait of Volcher Coiter, act. 41 (painted in r 575 byan
famous calculation about the circulation of the blood.
unknown master)


Freind's view that the matemnl and foetal circulations were continu-
Portrait of Sir Kenelm Digby (from the painting by
ous was derived from the experiments of Rayger and Gayant, who had
Oomelius Jansen, ca. 1650)
injected a blue dye into the foetal circulation and found it again in the
maternal. Vl/orlt of this kind had begun as far back as about I555, when
apparently Axuatus Lusitanus had made similar observations on a
woman, and in the Cracolfia of J. F. I-lertodt, published in 1671,
where under the heading “An erocus foetum tingat in utero ?" we find
a description of the public dissection of a pregnant dog to which this
dye had been given in the diet. The embryos were markedly yellow.


' Cf. the interesting recent resume of W. schopfer.
Portrait of Sir Thomas Browne and his wife,
Dorothy (ca. x650)


' Biographical detail: of Freind in an essay by Greenwood.
IX Zeus liberating living beings from an egg (the frontispieoe


' Muller-lies: has written I monognph on the development of knowledge on
of William Harvey’s book on the Generation of Animal:,
menstruation from the sixteenth century ouwuds.
1 6 51) ‘


X Illustrations from “falter Needl1am’s De Fonnatn Foetu of 1667


PLATES


TABLE I
Illustrations from Maipighfs 1): 01:0 Incubate of 167:
shgwmg the early stages of the development of the
°h“'-k facing page 166


I. That the embryo was nourished directly by menstrual blood.
Illustrations from 77:: auiou: and accurate ab.mvan'an: 0/
Mr Stephen Lorene-t'm' of Florence on the Dirxrrtioru qf


Beckher, 1633.
the Cramp-Fish ,9,
Metamorphosis in Buddhist iconography; statues in the


Plempxus, 1644.. Plcmpius did not deny that the umbilical cord
Sleeping Buddha Temple at Suchaw (Chin-dffian),
was functional, but insisted that the blood passing through it
was menstrual. In 1651 Harvey’: vmrk was published.


smnemfi. 1554» F. Sylvius, I680.
Kansu province. China ,7;
Portrait at‘ Antan van Leeuwmhoek by Job.


Seger, 1660. Cyprizmus, 1700.
Vex-kalje (1686) :7:
Portrait of Robert Boyle (a. 1690) 176
A tel-atoms with well-formed teeth and hair (from Robert


van Linde, 1672.
Plat‘: Natural Hi:-tmy afS!ufl'ord:h:'r:, 1686) 178
Illustrations from Antoine Maine-Jan‘: Obaematiovu


II. That the embryo was nourished through its mouth.
mr Infmnahbn du pcmlet (1722) 186
(:1) By the amniotic liquid.
De Réaumux-‘I Incubators (from De (‘art de faire (chm
(i) In addition to the umbilical blood.


Harvey, 1651. Linsing, 1701.
In paulm, 1749) 204
W. Nctdhaxn, x667. Pauli, 1717;.
de Gruf, 11977. Barthold, x717.
 
C. Bzrthulihus, 1679. S. Middlebezk, 1719.
van Diemerbroeck, 1685. Teichmeyer, 17x9.
 
Ordob, X697. Gibson, 1726.
D. Tauvry, 1700.
(ii) Alane; the umbilical blood being regarded as unneces-
mry or of minor importance.
 
Moellenbrcck, r672. P. Stalpartius, 1687.
Everardus, I685. Bierling, 1690.
 
Case, 1696. Case thought the embryo arose entirely
out of the amniotic liquid like a predpitate from 3
dm solution; see, p. 184.
 
Berger, 1702.
 
These writers assumed as their prindpal experi-
mental basis reports of embryos born without
umbilical cords, e.g. those of:
 
Rommelius, 1675 (in Velsch).
 
Valentinius, 1711.
 
(17) By the uterine milk or mu-um Iadzo-diylamm.
 
Mercklin, 1679. J. xvaldschmidt. 169:-
Drelincurtius, 1685. Tauvry. I694-
Bohnius, 1686. F73“? 1732'
Zacnhias, x688. D3011”: 177-4-
 
 
III. That the embryo was nourished through the umbilical cord only.
(a) By foetal blood (the circulations distinct).
 
Arantius, 1595. Snelle. 17o5.
Harvey, 1651. Falconnet, 1711.
W. Ncedham, 1667. F. Hoffman, 1718.
Ruysch, 1701. Monro, 1734.
 
It is to be noted that Bicrling, P. Stzipnrtius,
Berger, Barthold and Charleton, who supported the
discontinuity theory of the circulations, were all
upholders of the theory of foetal nourishment per 0:,
so that their reasons for doing so were not those on
account of which we agree with Hofimann and
 
Needham at the presmt time.
 
(b) By maternal blood (the circulations continuous).
Laurentiuz, 1600. Hamel, 17oo.
de Marchette, 1656. de Craan, 17o_-4.
Rallius, 1669. Lang. 1704.
Muraltus, 1672. van Home, 17o7.
Blasius, 1677. Freind, 1711.
Vcslingius, 1677.
 
de Méry, 1711. De Méry comhated Faloonnet's view
of the separate circulations. He said that he had not
himself tried Falconnetla experiments, but that
some students had, and could not repeat them.
 
Aubert, 1711. Narrative of a use in which the um-
bilical cord had not been tied at the maternal end
and the mother had nearly bled to death through it.
 
Nenterus, 1714. Wedel, 1717.
 
Bellinger, 1717. Bellinger believed that the maternal
blood was transformed by the embryonic thymus
gland into proper nourishment for itself, after which
it was secreted into the mouth by the salivary ducts
and so went to form meoonium without the necw
sity for dcglutination. Heister's comments on this
extraordinary theory are worth reading. Perhaps
Bellinger was indebted to Tauvry for his idea of the
importance of the thymus gland. Tauvry had dnwn
attention in 17m to its diminution after birth.
 
dc Smidt, 1718. Dianis, 172.1.
 
 
(continued overleaf)
181
A A 1-nsronr or EMDRYDLOGY '
 
new 1 (tonlirwed)
 
 
(t) by menstrual blood.
Plempius, 1644.
 
(J) By uterine milk.
 
Em, x687.
 
Cameraxius. I714. (Opfnfo concilialrixl)
 
F. Hotfmann, 1718.
 
(t) By the amniotic fluid.
Vicafills. 170°: Goelicke, 1723.
IV. That the embryo was nourished through pans in is skin,
 
Deusingiua, 1660. Stockhamer, X682.
 
Nitzsch, 1671.
This was suggested on the ground that in the earlier
stages of development there is no umbilical cord. In
1684 de_St Romain argued against it on the ground
1-bit: if I! Went true, the embryo would dissolve in
the amniotic liquid.
 
During this period also there were continued disputes about the
origin of the arru-riatic liquid. Van Diemerhmeck and Verheycn con-
sidered that it could not be the sweat of the embryo, for the embryo was
always much too small to account for it, and, moreover, du Tertre had
described cases where the secundincs had been formed with the mem-
branes but in the absence of the embryo. Dionis aflirmed that whatever
it was it could not be urine, for urine will not keep good for nine days,
afnrtian‘ not for nine months. Drelincurtius put forward a theory that
the embryo secreted it from its eyes and mouth by crying and salivating,
while Bohn and Blancard derived it from the foetal breasts. Lang,
Berger and de Gouey criticised this notion without bringing forward
anything constructive, and de Gouey was in his turn annihilated by
D. Hoffinzmn, who with Nenter and Kiinig supported the modem view,
namely, that it was a transudation from the maternal blood-vuseis in
the decidua. The question was complicated further by the alleged
discovery by Bidloo in 1685 of glands in the umbilical cord, and by
Vieussens in 1705 of glands on the amniotic membrane. J. M. Hoffmann
and Nicholas Hoboken supported the view that these were the impor-
tant structures. There the problem was left during the eighteenth
century, various writers supporting different opinions from tune to
time, and it is still not fully solved.
 
 
2. Growth and Differentiation; Stahl and Maitre-Jan
 
Very early in the eighteenth century (1708) there appeared a work by
G. E. Stahl, van Helrnont’s most famous follower, which struck the
keynote of the whole period. Stahl‘s Thearia Zlrledim Vera, divided as
it was into physiological and pathological sections, belonged in essence
to the a prion‘ school of Descarts and Gassendi. It differed from them
profoundly, however; for instead of trying to explain all biological
phenomena, including embryonic development, from mechanical first
principles, it started out from first principles of a vitalistic order, and,
having combined all the arthaei into one informing soul, it sought to
show that the facts could be convincingly explained on this basis.
The spiritual kinship of Stahl with Desmrtes and Gassendi is due to
an atmosphere which can only be called doctrinaire, and which was
common to them all. Like the methodist school of Hellenistic medicine,
they subordinated the data to a preconceived theory.
 
Stahl is also interesting in that he represents a trend of thought which
favoured what has been called "instantaneous generation followed by
metamorphosis.” Cole calls this the “precipitation” theory! but I cannot
altogether accept his account of it.
 
"Metamorphosis” was defined by Harvey (1653, pp. 222 if.) m the
bringing into being of a formed object from a mass of material previously
possessing no form; as opposed to “epigenesis," where assumption of
form and increase of mass proceed simultaneously.
 
Artificial productions are perfected two several waies; one, when the
artificer cuts and divide: the matter which is provided to his hands, and so
by paring away the superfluous parts doth leave an Image remaining behinde,
as the Statuary doth; the other, when the Potter forrnes the like Image of
Clay, by adding more stuff, or augmenting, and so fashioning it, so that at
one and the same time, he provides, prepars, fits, and applia his materials.
Harvey's "metamorphosis" we should now call "differentiation without
growth” and his “epigenesis” we should call "differentiation plus
growth." It should be carefully noted that to epigenesis Harvey does not
oppose “preiorrnation,” for he was writing some thirty years before
Malpighi’s unfortunate surnmer-time experiments, and Joseph de Aro-
matari's seed had not yet sprouted. "I’reformation” in modern temts
would correspond to "growth without differentiation," all the com-
plexity of the finished form being supposed to be present initially.
“Metamorphosis" in Harvey’s mind was nothing more nor less than the
Aristotelian blood-and-seed theory, and his description of :1 sculptor
making a statue out of a pre-existent mass can best be understood in
the light of Ruefl"s drawings (Fig. 10). Harvey opposed Fabricius pre-
 
‘ P. 205.
183
A HISTORY OF EMBRYOLOGY
 
cisely because the latter ventured to point to the chalazae in the hcn's
eg as the homologue of the mammalian blood-and-sccd,
 
But the relations between growth and difl'erentiation were still open
to different opinions in the eighteenth century and many believed rm:
'1" former was by far the more impomm ofthe two. If the di5erentia~
tion process was pushed far enough back in the life of the embryo the
distinction between epigenesis and preforrnation tended to disappear
We have already had (p. 167) in Croone an example of a prefer-rnationisti
Who bclieved illogimlly that “instantaneous generation” was followed
by "metamorphosis,” and Stahl is an example of an epigenesist who
thought that the governing artlmeur provided by the semen organised
whatever it found in the uterus into the form of the body, after which
there was pure growth and no further differentiation. The distinction
thus became almost academic.
 
After the mysterious organising process
the entire remaining business of generation [said Stahl‘] is taken up with the
fomzation of the body, which from the first rudiment, so to speak, is nothing
afterwards but nutrition; namely, such as is carried on from this time up to
old age; while in the body what is once completely shaped is not merely
preserved by a perpetual supply, nor, if it pen-hanoe fails, is it merely rebuilt,
but in fact it continually grows until it is completely formed in all its pans.
A perpetual assimilation everywhere accompanies apposition, or rather the
apposition itself is set up immediately in such order and situation that
assimilation is brought about and exists bemuse of that very position. . . .
That Principle which unfolds its activity primarily in the brain and the nerves
prmides over the formation of the body, and those parts which constitute the
only immediate instrument of in action: being formed first of all, make it
probable for that reason that something ought to, or at least an, be provided
by themselves alone. . . . As for how the blood is generated, it’ we are to be-
lieve the theories commonly held today, it is thought to be born from a
spontaneormalgllidhtg-together and chance meeting of particles uniting them-
selves mutu y.
 
Thus:
 
(A) EPIC!-‘N515 E Diflerentiation + Growth Harvey
 
(B) PRErDR.\tA‘I10. Growth alone Malpighi. Sw:rnmcrdzm.€t€-
(C) luzrluuoxrflosts Differentiation alone Aristotle, Fabriciu:
 
(D) PRECIPITATION E (A) in the very early Sub]. C359
 
stages, followed by (B)
(E) (B) in the very early stages, followed Croone
 
 
by (C)
(F) (C) in the very early stagm, followed Butfon
‘by (B)
Thewid. on 425 S-
 
134
EMBRYOLOGY IN THE EXGHTEENTH CENTURY
 
In 1722 Antoine Maitre-]an published his book on the embryology
of the chick, the only one on this subject between Malpighi and Heller.
It was an admirable treatise, illustrated with many drawings which,
though not very beautiful, were as accurate as could be expected at the
time. Perhaps its most remarkable characteristic is its almost complete
freedom from all theory——Maitre-Jan says hardly a word about genera-
tion in general, and is far from putting forward a "system" in the usual
eighteenth-century manner. He contents himself with the recital of the
known facts including those added by his own observations. He gives
no references, and writes in an extremely modem and unaffected style.
 
The only traces of theoretical presupposition which can be found in
him are Cartesian, for he speaks of the activity of fen-nents in blood-
fomzation. He is an epigenesist, and long before Brooks, he gives the
right explanation of Malpighi’s error, afiirming that the hot Italian
summer was responsible for some development in Malpighi’s eggs be-
fore Malpighi examined them. Although Maitre-Jan’s book must have
been accessible both to Buffon and Haller, they perpetuated Malpighi’s
mistake till nearly the end of the oentury.
 
In technique, Maitre-Ian was pre-eminent. He was the first embry-
ologist to make practical use of Boyle’s suggestion regarding “distilled
spirits of vinegar" for hardening the embryo so that it could be better
dissected.‘ He also used “weak spirits of vitriol"; after treating blaste-
derms with it, he said, “I saw with pleasure an infinity of little capillary
vessels which had not appeared to be there before” (see Plate XVII,
facing page r86). He made a few chemical experiments also, noting that
vinegar would coagulate egg-white, and estimating quantitatively the
difference in oil-content of different yolks—though for this he gives no
figures.
 
His theory he relegated to an appendix entitled Objettimz: :10‘ la
génératian dc: animaux par de petilr em‘. There were sixteen of them,
but the most cogent one was that, as little worms had been found under
the microscope in pond-water, vinegar and all kinds of liquids, there
was no reason to suppose that those in the semen were in any essential
way connected with generation. For his time, this argument was an
 
‘ His Ietuzl words are as follows:
 
p. nu. "Si l'on verse dan: cet zzuf (70 hours) nu dans un autre de pareil tems de
oouvée, du vinaigre distillé, on verrn tn peu de tam: le fetus blanehir ex devenir plus
solid: at xi opnque, qu'on no | uroit plus dixtinguer nu lntvers lea vésiculu du cczur,
m mam: lea vusseaux qui y s utissent, hon celui qui régne le lung du Carpl."
 
p. 148. :']e du-ni enfin que quoiqu‘on n: décourre quhssez ebseurément ls plflpnt
ties pnnapales parties intérieureu, hon le ctzur ct quelque: vnisseaux (144 houn),
I cause d:_ lcur trop dc moluse et de la viseoaité de tout le fetus, elles ne laisxent pas
que d'A_vmr sen quelque farme, comm: on 1: oonnolrn en fnisanr infuser le ftztux dun
le vinugre dxsulé."
 
:85
A HISTORY OF EMBRYOLOGY
 
excellent one, and was .
exiebrimen: which hflli)xX’1el)[ity‘:;:I?):leer:n!1!l]:§:VE3S:;I;l‘1)c §:oIr:1!1:)of filtration
out I 's time ' . ‘ _
blood, the fwamm ézgfélisiiilrgeezgtrgvexrsrygter the circulation of
“"1 d° M57 “"5 mgflgtd in a pnlcmi<?ori this stil7)?c:i° I7.i‘:'1,Tauwy
also corrwponded with Duvemey, snvcstm and ear: . e latter
“'°V°’5}’ “'hi°l\ refills that of Laurcntius and Petreus a hunrirn L: can.
before. Nichnlls wrote later on the same sulfect Daniel Ta e yyurs
i"‘°’°51i“8. h°W€V€1'. for other reasons. He  an epi  Wag
 
“"°‘° Vi8°'°“5lY agtinst the view that the soul constricted diuiinn
cmbryvgeny a suitable home for itself. 3
 
Nine years later two books appeared which form very notable land-
in the history of embryology. One was Martin Schurig’s Embryo-
IUEW. and the other the Elementa Clxemiae of Hermann Boerhaave
 
Th: {°m‘"v h°W¢V€|'. gave to the world no new experiment‘; of
°b5C1'V3"'0fl5; it Was the first of What we should now call the typical
“'°"i°“"' kmd °f Plllflimtion. Schurig saw that he was living at the end
of :1 great scientific movement following the Renaissance. and set him.
self accordingly for many years to Compile large treatises on specific
physiological subjects, taking care to give all references with meticulous
accul'-“»'}'- and to omit no work, significant or insignificant. His Spam.
tologfa was the first to appear (in 1720), and it was followed in 1723 by
Sialologia (on the saliva), Chylologia (1725), Muliebria (1729), Parxlmm.
1031}! (1719). Gyflllttnlogia (1731) and Hazmatologia (1744). His Embryo-
Iogm was the last but one of the series. In it he treated compendiously
of all the theories which had been advanced about embryology during
the immediately preceding two centuries, and his chapters on foetal
nutrition and foetal rcspiration throwa flood of light on the "intellec-
tual climate" in which Harvey and Mayow worked. Schurig‘s biblio-
graphy is a very striking part of his book, extending to sixteen pages,
and inlfll-‘l.‘l:.i-llg five hundred and sixty references, it was the first attempt
of its ‘ .
 
3. Chemical and Quantitative Approaches to the Origin of
Organisation; Boerhaave, Hamberger and Mazin
Hermann Boerhaave was a more prominent figure, a professor at
Leiden for many years, and renowned for his cncyclopaedic learning
on all subjects remotely connected with medicine. His Elemmta
C/zhniae, which became the standard chemical book of the whole
 
period, demonstrates throughout the exceedingly wide outlook of its
 
and contains in the second vmlume what must be regarded as the
 
author,
I reproduce here the
 
first detailed account of chemical embryology.
186
PLATE XVII
 
llluxlralmmfram Antoine fllailn-]n1|‘x Obsenutions sur la Iormznon du poulcl, oh la: dnen
changtmms qui arriu-nx i l’<:ufi mcsun: qu'|l 2:: com é, mm exacxemem cxpliqufi
ct rcprésnntéa en F: ns D‘IIour_v. Parix, 172:.
 
A. Drlvungs olcmbryos of rum. zoc»15o hours inzubznon.
 
n. The am dnlung u! lh: fillosines on uh: inlumr of the yoli-sac.
 
tp. 2,2) “um: pike .1. 1. “mad: membrane de_1:-me sur 1. ...,msc.e mttnmre at hqurlle on wit ..x..,.=un
nnglu d: m penis vusiuux cmnrnllées dc dnusn grlndeurs ti dxlhrement amass" L179 hrs 1
c. Photaznph at the .-.11:-mm. from Rcrnoui. They puy m impomm pan m mg Ibsotpllnn n( the nu.
 
 
um
EMERYCILOGY IN THE EIGHTEENTH CENTURY
 
relevant passages in full because of their great interest. It will be noted
that they are cast in the form of lecture addresses, as if they had been
taken down direct from the lectures of the professor, a fact which gives
them a peculiar charm when it is remembered how many great men
must have listened to them, among them Albrecht Von Heller and Julien
de la Mettrie. In considering what follows, it should be noted that Boer-
haave's interest is biological all the time, and that he does not trmt the
liquids of the eg. as nearly all the chemists before him had done, as
substances of curious properties indeed, but quite remote from any
question relating to the development of the embryo. Another interesting
point is that he deals only with the white, and hardly mentions the yolk;
this is perhaps to be explained by the Aristotelian theory‘ that the
embryo was formed out of the white, and only nourished by the yolk
(ex albofieri, ex luteo nutn'n'), a theory which was still alive, in spite of
Harvey, in the first half of the eighteenth century. If this was what was
at the bottom of Boerhaave's mind, then it is obvious that the egg-white
would be to him the liquid inhabited more particularly by the plastic
force. This, then, is what he has to say about the biochemistry of the
 
egg.
 
0]. Chem. in Am'nmIz'a. (Processus log.) The albumen of A fmh egg it not
add, nor alluzline, nor doe: it contain afzrmenled .tj:x'n'l. The white of a fresh
egg, separated from the shell, the membranes and the yolk, I enclose in clean
glass vessels, and into each of these I pour different acids, and shake them up,
mixing them, and no sign of ebullitiou appears however I treat them. Thero-
fore I lay these vessels aside. Now in these other two vessels I have two fresh
portions of albumen, and I mix with them in one case alkaline salt and in the
other volatile alkali. You see they are quiet without any sign of effervescenoe.
Now behold a remarkable thing, in this tall cylindrical vessel is half an ounce
of the albumen of an egg and two drains of spirits of nitre, in this other vessel
is half an ounce of egg-white, together with four and a half ounces of oil of
tartar per deliquium both heated up to 92 degrees. Pray observe and behold,
with one movement I pour the alkaline albumen into the acid albumen, with
what fury they boil up, into what space they rarefy the mass, so that they
stream out of the vessel although it is ten pints in size (decupli eapace). They
have scarcely changed their oolour. But when the eifcrvescenoe has abated how
suddenly they return to the limits of space occupied before. But now if more
egg-white is heated to 100 degrees in a retort (nu-urbihz) an insipid water
containing no spirit is given oil’. If egg-white is applied to the nalted eye or
naked nerve it does not give the smallest sense of pain, and scarcely affects the
smell; nothing more inert and more insipid can be put on the tongue. It
appears muoous and viscid to the touch, not at all penetrable. Hence in the
fresh white of an egg there is no alkali or acid, or both together. It is indeed
 
‘ Iliriarrh, 56:‘ :5.
187
A HISTORY or BMBRYOLOGY
 
3 ‘hick, Slick ‘. incfl. and insi id li uor, e ‘ ' ‘ -
heat of 93 degrees within the sgaee oci 2t dz):  ;::.i,'§§:,'§,ff,§
egg from a tiny mass hardly weighing a hundredth of a grain into the perfect
b°‘:1Y Qf 311 31151113]. Weighing an ounce or more. We have learnt therefore of
a liquid distinct from all others, from which by inscruublc mum mm,
membranes, vessels, entrails, muscles, bones, mrfilaggg’ and 311 lb, cum:
parts, tendons, ligaments, the beak, the claws, the feathers, and all the humaurs
can be produced-and yet in this liquid we find softness inertia absence of
Mid. alkali, and spirit, and no tendency to eflervesce. Indeled if there were the
slightest efl'ervescenee in it, it would certainly break the eggshell; therefore ne
see from how slow and inactive a mass all the solid and fluid parts of the
chick are constructed. And yet this iself is rendered absolutely useless for
forming the chick by greater best. It smrcely bears no degrees with good
effect but at a la: temperature never brings forth a chiclr, lorunder 80 degrees
will not sutfioe. But by a heat kept between these limits, there is brought
about so marvellous an attenuation of the mucous inactivity that it can exhale
a great part through the shell of the egg and the two membranes, the yolk
and chalazae alone remaining slang with the amniotic sac. For the yolk, the
uterine placenta of the chick, takes little part in the nourishment. Meanwhile
Malpigbius has shown that this albumen is not a liquid of a homogeneous
kind, as the blood-serum flowing through the vital vessels is, but that it is a
structure composed of numerous membrane-like and distinct small sacculcs,
filled with a liquid of their own, in the same way as in the vitreous humour of
 
e eye.
 
(Prooesus xxx.) Eajblonzlicm of the egg-white u-ill: almlrol. In this trans-
parent vessel is the albumen of an egg, and into it, as you perceive, I gently
pour the purest alcohol, so that it descends down the sides of the vessel and
reaches the albumen. I do this deliberately and with such solidtude that you
may see the surface of the albumen which, touching the alcohol, holds it up,
being immediately coagulated, while the lower part remains liquid and trans-
parent. As I now gently shake them together, it appears evident that wheret er
the alcohol touches the albumen a onncretian is formed. Behold now, while
I shake them up thoroughly together, all the egg-white is coagulated. If
alcohol previously warmed is employed in this experiment, the same result
is brought about but more rapidly. It appears therefore that the purest
vegetable spirits immediately coagulate the plastic and nutrient material.
 
(Prooessus :12.) Thefresh albumen afar: egg is broken up by di'm'!hzti0n.These
{rah egg have been cooked in pure water till they become hard. I now take
the shining white sepanting off all the other things and break it up into small
pieces. I put these, as you see, into a clean glass retort (nu-mbiln) and I duly
cover it by fitting an an alembic and add a receiver. By the rules of the
[chemical] art I place the whole retort in I bath of water and I apply to it
successive degrees of fire until the u hole bath is boiling. No vnporous strealzs
(mine) of spirits are given off but simple water  detvy drops and this in
incredible quantity, more than nine-tenths. I eonunue so with patience until
by the heat of boiling water no more drops of thn humour are given 05.
 
188
EMBRYOLOGY IN THE EIGHTEENTH CENTURY
 
Then this water shows no trace of oil, salt, or spirit; it is perfectly transparent
and tasteless, except that it eventually grows rather sour. It is odourless, save
that towards the end it gives ofi a slight smell of burning. It shows absolutely
no sign of the presence of any alkali, when I test it in every way, as you can
see for yourselves; nor does it reveal any trace of acid, when tried how you
will. Here you see pounds of this water, but in the bottom of the now open
retort see, I beg of you, how little substance remains. Behold, there are
fragments contracted into a very small space in comparison with the former
quantity. They are endowed with a golden oolour especially where they have
touched the glass, but yet they are transparent after the manner of coloured
glass. When I take them out I find them very light, very hard, quite fragile,
and breaking apart with a crack, smelling slightly of empyreuma, with a taste
rather bitter from the fire, and without any flavour of alkali or acid. This is
the first part of the analysis. Now I take these remaining fragments in a glass
retort (retorlam) in such a way that two-thirds remain over. I put the retort
into a stove of sand, first arranging a large receiver. Then thoroughly luting
all the joints I distil by successive grades of fire and finally by the highst
which I all supprrstianis. There ascends a spirit, running in streaks [striatim]
fat and oily, and at the same time, volatile salts of solid form everywhere on
the walls of the vessel, rather plentiful in proportion to the dried fragments
but small in proportion to the whole albumen before the water had been
removed from it. Finally an oil appears besides the light golden material
mixed with the first, black, think, and pitchy. When by the extreme force of
the fire this oil is finally driven forth, then the earth in the bottom, closely
united with its most tenacious oil, swells up and is rarefied and rises right up
to the neck of the retort so that had the retort been overfull it would have
entered into the neck and clogged it up even causing it to burst with danger
to the bystanders. The operation is to be continued till no more comes out.
That first spirit, oily and fatty, is clearly alkaline by every test, as you may tell
from the way it effervescm when acid is poured on it. If we rectify it we
resolve it into an alkaline volatile salt, an oil, and inert foetid water. The salt
fixed to the walls is completely alkaline, sharp, fiery, oily, and volatile; and
the final oil is specially sharp, caustic, and foetid. The black earth which
remains in the retort is shiny, light, thin, and fragile, foetid from the final
empyreurnatic oil, and soft because of it. If then it is burnt on an open fire,
it ‘mrres a ‘i':'t‘t're ‘i-med ta-nh 1v‘rnt.’n is ‘wloote,  , tasteltn, and odumitss,
from which scarcely any salt can be extracted, but only a very heavy dusty
powder (poIIx'nem).‘
 
(Processus H3.) Tlrefrerh albumen :1] an Egg zcillputrefy. Sound eggs kept
at 7o“ for some days will become foetid and stink. . . . \Ve have learnt then that
this is the nature of the material which will shortly be changed into the struc-
ture, form, and all the parts of the animal body. Repose and a certain degree
of heat produce that effect in that material. We observe therefore the spon-
taneous eorruption and change of the material, ahd what is extremely re-
markable, if an impregnated egg is warmed in an oven (in hypocaurrir) to a
 
‘ Cl. the dry distillation of egg-white by Pietet 5: Crime: in 1919.
189
A HISTORY OF EMBRYOLOGY
 
heat of g: degrees it employs these attenuated parts changed by such a heat
*0 ߢ_"1HSl1. Incrensc._nnd complete the chick for 2x days. But in this chick
nothing alkaline, foettd, orputnd L! found, henoe observe, 0 doctors (rnedt'¢1')
the remarkable manifestations of Nature—by repose and a certain degree of
heat a thick substance becomes thin, a viscous substance becomes liquid, an
odourlex substance becomes toetid, an insipid substance becomes sour and
extremely acrid and bitter to the taste, a soothing substance beoomea caustic,
a non-alkali become: alkaline, a latent oil becomes sweet and putrid. Let these
results be compared with the observations of Llarcellus Malpighius on the
incubated egg, and we shall observe things which shall surprise us. I took
are to investigate only the albumen of the egg Erst of all, separating the other
parts off where possible, for the albumen alone forms the whole of the
 
material which proceeds to feed (in pabulum) the embryo. The other constit-
uents of the egg only assist in changing the albumen, so that when it is
 
changed, it may be applied to farming the structure of the chitin
 
Boetl1aave's treatment of thae subjects has only to be compared with
that of Joachim Beecher, who wrote in 1703, to show how thoroughly
modern in outlook it is. Beccher's Pllyrita Sublerranea contains a whole
section devoted to the growth of the embryo, but it is extremely con-
fused and very alchemiml in its details.‘ The advance made in the thirty
years between Beecher and Boerhaave was immense, but, if the bio-
chemistry of development advanced so fast, its biophysics was not far
behind, as is shown by the work of G. E. Hamberger and I. B. Mazin.
 
H:trnl>erger’s most important contributions, contained in his Plryn'o-
lagia Media: of x751, were his quantitative observations on the water-
content of the embryo and its growth-rate, in which he had no fore-
runners. Hamberger showed
 
that there are much less solid pans in the foetus than in the adult. The
cortiml substance of the brain of an embryo loses 8694 parts in ro,ooo on
drying but in the adult it only loses 8096 and that of the cerebellum from 8t
parts is reduced to 12. The maxillary glands of the embryo lose out of ro,ooo
parts 3459. the liver 8047, the pancreas 7863, the arteries 8278 and even the
cartilage: lose four-fifths of their weight decreasing from xo,ooo to Buy}.
 
The corresponding figures for the adult were: liver 7r92, and heart
7836. These figures do not widely difler from those obtained In recent
Mazin published his Conjecture: physiro-medico-12;,-drottoiicae d:
Respiratione Foetu: in 1737 and his Tradatur Zlleniico-meelxonxta in 174-3-
In the first of these works Mazin supports “hat is rsserrtially Mayow s
theory of embryonic respiration, without, however, mentioning hlayow
more than once. It had not been popular since 1700, though Prtcarm
1 BL :, mt. iv, ch. 4. p. 1111, “De mixfione
 
I90
EMBRYOLOGY XN THE EIGHTEENTH CENTURY
 
had defended it. Mazin put the liquids of eggs under an air-pump, and
observing that air could be extracted from them aflirmed that the air was
hidden in them and that the embryo could therefore respire. He spoke of
“aerial particles" in the amniotic liquid, and discussed the respiration
of fishes in connection with this. The specific gravity of the embryo also
interested him, and he did a great deal of calculation and experiment on
it. His most interesting passage, perhaps, is that in which he mentions
the “eolipilc" of the Alexandrians, the primitive fonn of the steam-
engine, and says thatjust as the heat of the fire makes the water boil, so
the heat of the viscera makes the amniotic liquid boil, giving ofl’
respirable vapours. The time-relations of this analogy are interesting,
for by I712 Thomas Newcomen had succeeded in making a stearn~
engine which worked with considerable precision, and the question of
steam-power was widely dismissed. Possibly Mazin was acquainted with
the Marquis of VVorcester’s Century of the Name: and Standing: of
Inventions, which had been published in 1663, and which had contained
an aeolipile or “water-commanding machine.” England was the centre
of this movement and other countries employed Englishmen as engi-
neers; Humphrey Potter, for instance, erected astearn—engine for pump-
ing«at a Hungarian mine in 1720.
 
As for the discovery of oxygen, it was near at hand, and Scheele in
1773 and Priestley in 1774 were soon to supply the knowledge without
which Mazin could not proceed further.
 
In his second book, Mazin reported many quantitative observations
on the specific gravity of the embryo. He found that it diminished as
development proceeded, being to the amniotic liquid as 282 to 274. in
the fourth month and as 504 to 494 in the fifth month. His work on this
subject was continued by Joseph Onymos, whose De Natura Foetur
appeared in r745. .
 
R. I. Raisin also contributed to this wave of precise measurement in
embryology. His dissertation of 1753 took account of the difference
between the pulse rates of infants and adults, and contained an arith-
metical argument about the prenatal secretion of the foetal kidneys.
But it also gave a list of the relative weights of organs, showing that
some decreased and others increased relatively to the weight of the body
as a whole. Thus the brain was one-tenth part of the body in the foetus
and onc~twenty-fifth part in the adult (see Table II, overleaf).
 
It was the first mention of heterauxctic growth,‘ save for the isolated
observations of Leonardo (see p. 98).
 
About this time. we get occasional references to the obscure mechan-
isms oontrolling animal growth. Although the brilliant speculations of
Marci (see p. 81) had long been forgotten, some writers, such as
 
r9r
A HISTORY OF EMERYOLOGY
 
TAB LE I I
 
F tut I
Cenbnm . . . . . r]'ro 11/12?”
Pulmo . . . . . r/66 r/r7
77‘J>mu: . . . . r/37.4 1/4560
90' - . . . I/189 r/1 r4
HfP'"' - . . . 1/23 1/21
I-"" - < - - - 1/324 1/175
Pancreas . . . 1/907 1/445
Vmtntulur vacmu . . . 1/767 1/212
Rene: . . . . . r/154 I/136
Surrenaler glarzdulae . . I/324 1/3040
 
Iames Parsons, were groping about for the rnorphogenetic controls.
In his Philoroplrital Observation: of 1752 Parsons had a good deal to say
about “primary" and "subordinate organisations,” notions which have
a certain resemblanoe to the field theories of modern embryology, for a
short account of which the article of Waddington may be consulted.
 
Parsons said of his organisations:
 
There can be no more natural Way of answering a Question proposed bya
Gentleman of Penetration in Philosophical Knowledge, which is Why do not
Animals and Vegetables grow on without End? Why do not Seeds, when
they are perfectly form'd, grow on in their Poet, Husks, or other Receptacles?
Because, says he, when a Body has once begun to grow, the same Propensity
for growing on ought still to continue, and, the Particles of Matter increasing
too, it ought not to cease.
 
The answer of Parsons was that the organisation comes to "its full
Power of Distention, so far as is consistent with its natural Form,"
after which further nourishment becomes useless and the structures all
Parsons was on the verge of a field theory, for he gave much
consideration to the regeneration arperiments carried out by Trembley
and others on fresh-water coelenterates (cf. Baker). But he did not
develop his idea far enough to escape the objection that the "organisa~
tions" were mere abstract sirnulacra of the visible forms of the animals
and plans themselves. For the test, he was a convinced ovist, accepting
Nuck's experiment (see p. 163) in the wrong sense (contrast with
Mamuet, see comment on p. 209), and desirous of explaining all genera-
tion as budding or "propagation." Like Galen long before him. ‘he
conducted a lively polemic against all formulations of the C13 plartztd,
but in favour, unfortunately, of the direct action of Cyod. In the succes-
sive action of his primary and seoondary 01’83“i53“°”5- h°‘""""" “°
 
I The classical modem treatment of the relative growth-nit! of mm cf °rz=fl"=m'
is of umne that of Huxley.
 
‘ P17. 94 E.
 
. 192
EMBRYOLOGY IN THE EIGHTEENTH CENTURY
 
approach rather closely the modern conception of a succession of
organisers or inductors in development (see Aristotle's ideas on this,
p. 48 passim, and the references to modern embryology there given).
 
These writers, together with Haller himself and J. C. Hefiter, who
handled the problem of embryonic growth rate, contribute to one of
the best, because most quantitative, aspects of eighteenth-century
embryology.
 
4. Albrecht van Ballet and the Rise of Techniques
 
Boerhnave‘s greatest pupil was Albrecht Von I-laller} Like Oliver
Wendell Holmes at Harvard, Haller occupied a “settee" rather than a
"chai.:” at Gfittingcn, and taught not only physiology but also medicine
and surgery, botany, anatomy and phamtacology as well. Nor did he
merely deal with so many subjects superficially; in each case he published
what amounted to the best and most complete text-book up to then
written. Haller was made professor in 1736, and for many years worked
at Gottingen, devoting much of his time to embzyological researches,
which, with those of his opponent Wolfi, stand out as the greatest
between Malpighi and von Beer. In 1750 he published a series of disser-
tations and short papers on all kinds of physiological subjects, which
would have been the direct ancestors of the modem compilations by
groups of experts, had they been more systematically arranged. The
volume on generation repays some study. The contributions relevant
to the present d'scussion. had been written at various times during the
previous seventy years, and may be summarised selectively as follows:
 
IV. Christopher Sturmius, Deplrmtarum animalimnque Generation: (first
published 1687). In this paper Sturrnius argues on behalf of the pre-
forrnation theory, "which in our times does not lack supporters,”
quoting Perrault, Harvey and Desmrtes. He contents himself with
countering arguments which had been urged against it, as, (a) spon-
taneous generation, (6) annual recurrence of plants, (c) insect meta-
morphosis, (-1) generation without copulation.
 
V. Rudolf Jacob Carnerai-ius,' Specimen Eajverimentomm pIr_y.rt'o-
lagim-thrrapeuiicomm circa Gmerationevn hominir el animalium. The
most interesting thing about this is that Camerarius mentions the
observations of D. Seiller, a sculptor, who had ascertained that the
body is five times the size of the head in the embryo but seven and a
half times the size afit in the adult. This is in the direct line between
Leonardo and Scammon.
 
XV. Philip Gravel, D: Superfzlalione (first published 1738).
 
‘ See d'I1sx3'.
' Afterward: furious as the discoverer of Izxualiry in planu.
 
n.L—~r3 I93
A rrrsrorw or-' rmaxronoay
 
XVIII. Adam Brettdel, De Embrymrz in 0911/0 mile amteplum pm:-e.u':tan1:
(first published x703). Breudel "stands for :1-.¢ Gmfim hyPo_
thesis. Unfortunately, he was also_a preforrnstionist and believed
that every limb, organ and function qisted not potentially hut
alaséueally in the urtfemlised eggbefore its passage down the Fallopian
 
XXII. Camillus Faloonnet, Non at friui Szmguir Itzgzmm az;,,,,,,;g (am
 
V published 1711). This is the first of the French u)mfibufi¢m_q M m,
 
book; they are all very markedly shorter than the Gemnn ants and
 
much less heavily omnmented with irrelevant quotations. 1-‘alommgg
 
is concerned to prove that the maternal and foetal circulations are
 
separate, and he describes in an admimbly eoncise manner ztn etperi.
 
ment in whid: he bled a female dog to death, after which, opening the
 
uterus, he discovered that the embryonic blood-vessels were full of
 
blood although those of the mother had none in at all. Ar-antius was
therefore justified. Falconner was soon confirmed by Nunn.
 
XXIII. Jean de Dir.-st’s Sui Sargguinix rolur gag’/es: Fetus at [first rzublished
173 5) was written to prove a similar point. He refers to the experi-
ment of Falcounet and the injections of F. Hofimann, and criticises
Cawper’a experiment in which mercury had been injected into the
umbilical vessels and found in the matemal circulation. on the
grounds that mercury is so “tenuous and voluble" that it might
pass where blood could not pass normally. He also objects to the
view that the foetus is nourished by the amniotic liquid.
 
XXIV. Francis David Herissant, Szmrrdinaeferui Pulmonir pnmfnnl aflicid,
:1 sanguine maltnw Fetus non alum (firs: published in X741). An
excellent paper in which the respiratory function of the placenta is
proved by the observation that the foetal blood-Vessel leading to the
placenta is always full of dark venous blood, while that leading away
from the placenta is light and arterial (flan‘dx'an‘ mrdnmque mlore, ul
iprzrnel obsemzd). Herissant adduces also the cases of acephalic
 
monsters, such as that of Brady, which could not poxibly have drunk
up any amniotic fluid, and yet were fully formed in all other respects.
He concludes that the urnbiliml cord serves for respiration and
nutrition.
 
XXV. After these three French workers, there is a great drop to Johnrtnes
Zeller, whose Injanlicidm mm abralt-it nee a tarlrna liberal Pubnmmn
Infantir in aqua nab:-idrnlia (first published 1691) is a long-winded
discussion of the floating lung test in forensic medicine. I-{is memory
deserves a word of ohloquy for his vigorous insrst.ence‘upon death
and torture for infanticide even during puerpenl insanity. Perhaps
it was Zeller who called forth the noble answer of de la Mettrie to this
inhumanity in his Man a fllaclxint.’
 
XXVI. Zeller’s De Vila humamz 2: F rm: pendent: (first published X692) is no
 
' For a detailed hisrorial amount of this tut and its unreliability, lee Knmmer.
‘94
EMBRYOLUGY IN THE EIGHT!-IENTH CENTURY
 
better, though at the time, perhaps because of its striking title, it was
famous. It deals with the ligation of the umbilical cord at birth.
 
Such were some of the typical papers printed by Haller in his 1750
collection. He retired from the Gottingen chair three years later, and in
1757 the first volume of his Elementa Phyrialagine was published,
probably the greatest text-book of physiology ever written. It appeared
only by slow degrees, so that it was not until 1766 that the embryo-
logical section was available. This volume contains a discussion of a
mass of literature, most of which had arisen during the preceding
twenty-five years, for although many of the names mentioned by Haller
occur also in Schurig, many are quite new.
 
Holler himself published in 1767 a volume of his collected papers on
embryology, most of which were concerned with the developing heart
of the chick, which he worked out very thoroughly, in collaboration
with Kuhlemann. Kuhlemann had already shown in the sheep what
Harvey had proved for the doe. But Haller was a convinced prefomia-
tionist, a fact which was largely due to his researches on the beds egg,
where he observed that the yolk had a much more intimate connection
with the embryo than had previously been supposed. Since the whole
yolk was part of the embryo, as it were, the preformation theory seemed
to him to fit the facts better than epigenesis.‘
 
Haller went further than Schurig in that he usually gave an opinion
of his own after summarising those of other people, but his views were
by no means always enlightened, and the atmosphere of Buflon is, on the
whole, more congenial to us than that of Haller. Haller, for example,
believed that the amniotic liquid had nutritious properties, and that the
nutrition of the embryo in mammalia was accomplished first of all per or
and afterwards per umbilimm. He denied that the placenta had any
respiratory function, and indeed his whole teaching on respiration was
retrograde. He mentions, however, an experiment of Nicolas Lemery's,
in which it had been found that indigo would penetrate the shell of a
developing hen's egg from the outside. Consequently. air might do so
too, and Vallisneri had shown that, if an egg was placed in boiled water
 
undcr an air-pump, the air inside would nrsh out through the shell and
appear in the form of bubbles.
 
I-laller was much more progressive in holding the origin of the
amniotic liquid (according to him a subject of extraordinary difficulty—-
rolurianem non promitlmn) to be a transudation from the maternal
blood-vessels. He followed Noorrwyek in asserting the separateness of
the maternal and foetal circulations in mammalia. He opposed the
 
‘ Sec pp. 2578-9.
195
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flu gvowlh In llnglh and weight of  berm III the (In: .
 
196
EMBRYOLOGY IN THE EIGHTEBNTH CENTURY
 
existence of eggs in Vivipara—“\Ve may conclude from all this,” he
said, "that the ovarian vesicles are not eggs and that they do not contain
the rudiments of the animal." But he accepted it in the restricted sense
that the embryonic membranes resembled an egg, thus:
 
If we call an egg a hollow membranous pocket full of a humour in which the
embryo swims, we may admit the opinion of the older authors who derive all
animals from eggs with the exception of the tiny simple animals of which we
have already spoken. It was in this sense that Aristotle and Empedoclcs before
him, said that even trees were oviparous. This has also been confirmed by the
experiments of Harvey on insects, fishes, birds, and quadrupeds.
 
Haller’s most original work was in connection with the growth-rate
of the embryo; here he struck out, for once, into entirely new country.
He made a beginning with the quantitative description of embryonic
growth, and one of his tables showing the changing lengths of the bones
is reproduced herewith (Fig. 21). He wrote:
 
The growth of the embryo in the uterus of the mother is almost unbeliev-
ably rapidr \Ve do not know what its size is at the moment of its formation,
but it is cenainly so small that it cannot be seen even with the aid of the best
microscopes, and it reaches i.n nine months the weight of ten or twelve
pounds. In order to clear up this speculation, let us examine the growth of the
chick in the egg. In this case again we are unable to measure its size at the
moment when the egg is put to incubate but it cannot be more than 1%.: in.
long, for if it were, it would be visible, and yet 25 days later it is 4 ins. long. Its
relation is therefore as 64 to 64 millions or r to r million. This growth takes
place in a singular manner; it is very rapid in the beginning and continually
diminishes in speed. The growth on the first day is from 1 to gr}, and what
Swammerdam calls a worm grows in one day from one twentieth or one
thirtieth of a grain to seven grains, i.e. it increases its weight by t4o to 24o
times. On the second day the growth of the chick is from r to 5, on the third
day, from x to not quite 4, on the fifth day from r to something less than 3.
Then from the sixth to the twelfth day, the growth each day is hardly from 4
to 5, and on the twentyiirst day it is about from 5 to 6. After the chick has
hatched, it grows each day for the first 40 days at an approximately constant
rate, from 29 to 21 on each day. The increase of the first twentyfuur hours is
therefore in relation to that of the lastns 5462 to 5 or r45 to r. Nowas the total
increase in weight in the egg is to that of the whole growth period (up to the
adult) as 2 to 24 ozs. all the post-embryonic growth is as 1 to xz, i.e. it is to the
growth of on: day alone early in incubation as 1 to 7§. . . . The growth of
man, like that of the chid-z, decreases in rapidity as it advances. Let us suppose
that a man, at the instant of conception. weighs a hundred-thousandth of a
grain and that a one-month-old embryo weighs 3:: grains; then the man will
have acquired in that time more than 3oo,ooo tima the weight that he had to
begin with. But it‘ a foetus of the second month weighs 3 ozs. as it approxi-
 
197
A HISTORY or EMBRYOLOGY
 
mately does, he will only now have a uired 8 times the mi ht e
 
beginning of the period. This is a ;l’qDdlgi0llS decrease in sgeedl alriildantl
end of the ninth month he will not weigh more than about ro5 o-zs. which is
not more than an average increase of 15 per montln A child three years old is
about half the size of an adult. It‘ then the adult weighs 225:: ozs. the three-
year old child only weighs 28i ozs. which is an eighth of the adult weight.
Now front birth to 3 years he will grow {mm ms to 281 or as 5 to 14, but in
the following 22 years he will only accumulate 2150 ozs. or eight times vthat
he had at 3 years. The growth of 3 man will therefore be in the first month of
intra-uterine lifeas r to 300,000, in the second as t to 48, in each ofthe others
as r to 15. In the first; years ofextra-uterine lifehis growth will hefrom :64.
to 281 and in the succeeding 22 years from 281 to 384., and the growth of the
first month to the last will he as 300,000 to -5%. or i36,8oo,ooo to 28, or
4,885,717 to 1. The whole growth ofrnan will consequently be as io8,ooo,ooo
 
to I.
 
In spite of the rather unfamiliar language in which these facts are
described, and the theory of the growth of the heart which Haller
subsequently put forth to explain them, they remain fundamental to
embryology. Their quantitative tone is indeed remarkably modern. In
my opinion, when all the voluminous writings of Haller are carefully
searched through, nothing more progressive and valuable than these
figures can be found. Heller and Hamberger stand thus between
Leonardo on the one hand and Minot and Brady on the other. That
they stood so much alone is only another indication of the extraordinary
reluctance with which the men of past generations assented to the truth
contained in Robert M:iyer’s immortal words, “Eine eirizige Zahl hat
mehr wahren und bleibenden Wert als eine kostbarc Bibliothelr van
Hypothusenl”
 
Of development as a whole, Haller spoke thus,
 
In the body of the animal therefore. no part it made before any other part,
but all are formed at the same time. If certain authors have said that the
animal begins to be formed by the backbone, by the brain, or-by the heart,
if Galen taught that it was the liver which wastirst formed,  others have
said that it was the belly and the head, or the spinal marrow with the bum,
adding that these parts malte others_in turn; I think that all these authors
only meant that the heart and the bfllfl or whatever organ it wrm, were nsibie
when none of the other pam yet were, and that certain parts of the embryoruc
 
body are well enough developed in the first few days to be seen While 015"!
are not so until the latter part of development; and others again not till after
birth such as the beard in man, the antlers in the stag, the breasts and the
second setof teeth. If Harvey thought he dscried an epigenetie developruent.
it was because he sawfirst I little cloud, then the rudunents of the head, with
 
I Iflrinne Sdtrijlm mi Bride, p. :26.
1 98
IZMBRYOLOGY IN THE BIG!-ITEIZNTII CENTURY
 
the eyes bigger than thewhole body, and little by little viscera being formed.
If one oompares his description with mine, one will see that his description
of the development of the deer corresponds exactly with mine uf the develop-
ment of the chick. If more than twenty years ago, before I had made many
observations upon eggs and the females of quadrupeds, I employed this
reasoning to prove that there is a great difference between the foetus and the
perfect animal, and if I said that in the animal at the moment of oonception
one does not find the same parts as in the perfect animal, I have realised
abundantly since then that all I said against preforrnntion really went to
support it.
 
The reasons for this change of opinion are not clearly given in Haller's
 
Writings, and Dareste concluded that it would always remain a mystery.
This mystery has, however, been almost completely elucidated by
 
F. J. Cole. In 1744 Haller was certainly an epigenesist,‘ in X758 un-
 
doubtedly a prefonnationist; in the intervening period he had made his
 
own embryological researches. How was it that they had the unfortunate
 
efiect of carrying him further from the truth rather than towards it?
In Cole's words:‘
 
The yolk, Haller asserts, is the continuation of the intestine of the embryo
chick. The inner membrane of the yolk is continuous with the inner mem-
brane of the intestine, and is thus identical with the inner membrane of the
gut generally and the skin and the ectoderm. The external membrane of the
yolk is an extension of the external membrane of the intestine, and is hence
continuous with the mesentery and peritoneum. The envelope which covers
the yolk during the last ten days of development is the skin of the foetus.
Therefore it is no absurdity to say that from the beginning, and before
fertilisation, the intestine of the foetus is no more than a small hernia of the
membrane of the yolk. Now if the yolk is continuous with the skin and in-
testine of the foetus it must be contemporaneous with it, and is truly a part of
the foetus. But the yolk was present in the abdomen of the hen, and was a
part of the hen, independently of any congress with the cock. Hence the
foetus, enclosed in the amnion, must have existed at the same time, though
invisible on account of its smallness and transparency.
 
It is not difficult, with the aid of Fig. 22, to follow Haller‘s argument. The
“inner membrane of the yolk" is the endoderm, which it is true does become
continuous with the skin and epidermis 11]!!! the gut cavity has been com-
pleted. The “cxtemal membrane of the yolk" is the splanchnic mcsoderm,
and the "envelope which covers the yolk during the last day: of incubation"
is the allanto—chorion, 's\ hich, however, is not the skin of the foetus. I-lallefs
procedure is typical of the time, in which observation and inference had only
the remotest relations with one another. For example, the statements "Now
if the yolk is continuous with the skin and the intestine of the foetus, it must
 
1 - . -
' gfneljhn not:g“on.l:.t'o;t'l!:zasa.vea Pnulcmonzx, vol. 5, pt. :1, pp. 497 fi’. (1744 ed.).
 
199
A msronr or IZMBRYOLOGY
 
_amu1 main...
 
 
Fir. 22. Dmgmn of the mmbmm o/ the rhiclt embryo, to tflurtmle I-IalIn'r
argument: (from F. 7. Cole).
 
be contemporaneous with it" and "the yolk must have arteries and veins as
without them it could not have been brought into existence" are pure
assumptions, and beg the very question he has set out to prove. If these
assumption: can be shown to be baseless, as they are, the whole argument
 
collapses.
 
In a word then, Haller confused the vitelline membrane with the
yolk-sac, and assumed a pn'z7n' that foldings, outgrnwths, etc., of cell-
layers could not take place, i.e. that epigcnetic processes did not exist.
Substantially the same argument as Haller’s had been used some twenty
years before (in 1722) by Maitre-Jan, according to Cole.
 
The zmbollzmznt aspect of prefomution presented no difficulties to
I-Ialler. Speaking of the generation of Volvax, he said,
 
It follows that the otary of an ancestress will contain not only her daughter
but also her granddaughter, her greatgr-anddaughter and her gre:Itgfeat-
granddaughter, and if it is once proved that an ovary can contain many
generations, there is no absurdity in saying that it contains them alL
 
200
EMBRYOLOGY m ‘rm: EIGHTEENTH CENTURY
The following passage is interesting.
 
We must proceed to say what is the efficient cause of the beautiful machine
which we call an animal. First of all let us not attribute it to chance, as Ofrai‘
would have us do, for although he pretends that all animals come from earth,
he is not attached to the ancient opinion, and nobody now believes what
Aelian says, namely that frogs are born from mud. . . . Vallisneri has found
the fathers and mothers of the little worms in galls, a quest of which Redi
despaired, and Redi‘ in his turn has made with exactitude and precision those
experiments which Bonannus, Triumphet, and Honoratus Faber had only
sketched out imperfectly. Moreover, no seed, no clover. . . . This was the
received opinion but in our century a proscribed notion has been revivified
and some great men have pretended that there are little animals which are
engendered by an equivocal generation without father and mother, and that
all the viscera and all the parts of these animals do not exist together, but that
the nobler parts are formed first by epigenesis and that then the others are
formed little by little afterwards.
 
This is an admirable illustration of how the ideas of spontaneous
 
generation and of epigenesis were bound up together. Haller goes on
to say:
 
M. Needham does not admit an equivocal generation but he does admit
epigenesis, and a corporeal non-intelligent force, which oonstruets a body
from a tiny little germ furnishing the necessary matter for it. He says that
there are only the primitive germs which were made at the original creation
and that germs organised like animals do by no means pre—exist, for if they
did, mole: ulninae, encysted tumours, and the like, could not come into
being.
 
Haller then goes on to describe Needham’s experiments with meat
broths, etc., and objects to his “system," largely on the ground that
“blind forces without any intelligence, could hardly be able to form
animals for ends foreseen and ready to take their places in the scheme
of beings." He considers that Needham's theories are completely dis-
proved by experiments such as those of Spallanzani, though, curiously
 
enough, he does not quote the latter author in this connection. I shall
return to this later  21 t). He continues,
 
Nobody has upheld epigenesis more than M. Wolff, who has undertaken
an examination to demonstrate that plants and animals are formed without
a mould out of matter by a certain constant force which he calls “essential"
(in his Tlieorfa Gmnalionil). . . . I have indeed seen many of the phenomena
which he describes, and it is certain that the heart seems to be formed out of
 
' is this not Iulien Offrxy de la Mtttrie? Hallcr had A habit of using Christian nazncv.
 
2.1:. Turbervdle {or J. T. Needham.
' See R. Cole.
 
201
A msronr or EMBRYOLOGY
 
n congealed humour and that the whole animal appears to have the same
consistency. But it does not follow that because this primitive glue which is
to take on the shape of the animal does not appear to possess ix; smmum and
all its parts, that it has not eiiectively got them. I haxe often given greater
solidity to this jelly by the use merely of spirits of wine and by this means I
saw that what had appeared to me to be a homogeneous jelly was carnpmed
of fibrm, vessels, and viscera. Now surely nobody will say that the vi:
mmlialir of the spirit of wine gave an organic structure to an unfcu-med
matter, on the contrary it is rather in the removal of transparency and the
accession of greater firmnuu to the extremities, as well as the making of:
more obvious boundary to the contour of a viseus that one could see the
structure of a cellular tissue. which was ready to he formed but which the
transparency had previously hidden and the wetness not allowed to be cir-
cumscribed by lines. . . . Finally, to cut a long story short, why does this ti:
crrentialir, which is one only, form always and in the same place: the parts of
an animal uhich are so ditferent, and always upon the same model, if in-
organic matter is susceptible of changes and is capable of taking all sorts of
forms? Why should the material coming from a hen always give rise to I
chicken, and that from a peacock give rise to a peacock? To these question:
no answer is given.
This was the case because Wolff was not a theorist, but rather an experi-
mentalist; his writings are marked by their abstention from the discus-
sion of speculative points. The above passage is very interesting. It
reminds us of the great difficulties with which the emhryologists of this
epoch had to contend. Serial section cutting was unknown, the staining
of thin layers and reconstruction were unheard of; even the hardening
of the soft embryonic tissues was only just discovered, as is indicated
by Haller above. Hertwig has excellently discussed‘ the advances in
embryological technique it hich took place during this and the {allowing
century. It is true that dyes were beginning to be used, as some instance
already given demonstrate, and as is seen from the use of madder in the
staining of bones, which began about this time, and was later much used
by the Hunters. Hertodt's Crocolagia is important in this connection.
Hertodt, by injecting saffron into the maternal circulation, found it
afterwards in the amniotic fluid,‘ and his experiment was cited by Heller
in support of that t.heory of the origin of the llq-llld.  the most im-
portant advance in technique was the progress In amfiaa-' tncuil-‘CM-‘L
The art, though lost throughorixt the Middle Ages and the seventeenth
century, was now to be revive . _
During this period much nor): was done on it. As far back as 1600,
de Serra had mentioned some experiments of this nature, but they were
not successful.‘ "The chicks." 11¢ Said» "“’"= USWHY 1”” d°r°”“°d’
 
1 And more recently Oppenheimer. ' Quzemo W. P- 273- ' m‘- "- ‘hr ‘-
 
201'.
I-ZMBRYOLOGY IN THE EIGHTEENTH CENTURY
 
defective or having too many legs, Wings, or heads, nature being
inimitable by art." Birch, in his History of the Royal Society, speaks simi-
larly.‘ “Sir Christopher Heydon [a relative of Digby's Sir Iohnij to-
gether with Drebell, long sincein the Minories batched several hundred
eggs but it had this efiect, that most of the chickens produced that \\ ay
\vere lame and defective in some part or other." Antonelli states that
similar trials were made at the court of the Grand-Duke Ferdinand II at
Florence about 1644, while Poggendorfi and Antinori relate that the
Accademia d. Cimento, inspired by Paolo del Buono, made trial of
artificial incubation between 1651 and 1667.
 
But the most famous of all the attempts to make artificial as successful
 
as natural incubation were those of de Réaumur, whose book De l‘ar1 de
faire [clove ler Pouletr of 1749 achieved a wide renown. He devotes many
chapters to a detailed description of incubators of very various kinds
(see Plate XVIII, facing page 204): but he nowhere gives any indiution
of his percentage hatch. It was probably low. He speaks also of the
“funestes eflets” of the vapours of the dung on the developing embryos,
without, however, furnishing any foundation for an exact teratology. In
the second volume he describes those experiments on the preservation
of eggs by varnish which caught the imagination of Maupertuis and were
held up to an immortal but by no means deserved ridicule by Voltaire
in his Akakia. For the details of this amusing but irrelevant issue, see
Miall and Lytton Strachey.
 
After de Réaumur, there were numerous continuations of the work
which he had started, in particular by Thévenot, La Boulaye, Nelli,
Porta and Cedemhieb-n. Much the most interesting of these was the
work of Beguelin, who attempted to incubate Qggs with part of the
shell removed so as to form a round window. He was not, however,
successful in the carrying out of this very modern idea. Probably the
most peculiar investigation made on developing eggs at this time was
that of Acbard, who is mentioned in a passage of Bonnet's.
 
M. de Réxurnur did not suspect in 1749 that some day one would try to
substitute the action of the electric fluid for his borrowed heat. This beautiful
invention was reserved for M. Achard of the Prussian Academy who excels as
an cxperimentalist. He has not so far succeeded in actually hatching a chick
by means of so new at process, but he has had one develop up to the eighth day,
when an unfortunate accident deranged his electrical apparatus.
 
Bonnet goes on to say that this substitution of electricity for heat
him hope that by electrical means an artificial fertilisation will one day
become possible.
‘ Vol. 3. p- 455-
2°23
A msronr or EMBRYOLOGY
 
References to these experiments and to those of - ~ ._
gators will be found in Heller. By the heginningmnofythlemxiiillezizecilth
Century :1 great mass of literature had developed on the subject, and it
possible to hatch out  more or less successfully from
 
, Oughfhe losses were stlll great. Early in the nineteenth
century Bonncrnaln and Iouard referred to the large number of mgnsms
produced, and in r8o9 Paris wrote,
 
_ During the period that I was at College, the late Sir Busick Hanrood. the
ingenious Professor of Anatomy in the University of Cambridge, frequently
attempted to develop; eggs by. the heat of his hotbed, but its only raised
monsters, a result which he attributed to the unsteady application of the heat.‘
 
5. Embryos and ‘Theologians
 
'I:his is the most convenient place to mention theological embryology
again. See pp. 22, 65-6, 75. Its place in the eighteenth century was small,
and in the nineteenth, with the general recognition that whatever the
soul might be it “as not a phenomenon, it altogether disappeared from
serious general discussion. F. E. Cangiamila's Embr;-alagia Sacra, how-
ever, ran through several editions between x7oo and I77 5. Cagniamila or
Cangi:unila' (Pammn. Ecrl. Can. 172:0]. at in tom Sic-il. Regan tantra
hatrttftam pm:-[totem Ingmlitore Pror:irla'aIi) deals very fully with the
time of animation, quoting a host of writers such as St Gelasius, St
Anselm, Hugh of St Victor and Pier) dclla Mirandola. His mind retains :1
quite mediaeval conformation, as the following curious passage illus-
trates: Quo! nonfazlus abortive: ex ignomnlia obrtetritmn rt nmmml exdpit
lafrfno, quorum zmima, :1’ Baptirmaie nan fmmiarelur, Damn in nztmuzm
ridntt. met demllius lumukmduml His instnletiorts fur the baptism of
 
I ormmam, p. 366. _
' Canglamlla, whose strnnge personality has recently been sympztlheualiy reviewed
by Hutchinson and hv Boldnm, deserves - lmle biognphienl notice. Born at Palen-no
in I702, he became Archprtesl at Glrgenti in 1731 and had the dsunctlon ofbzptinlng
the fiat infant delivered by Caesarean section in Sicily. He was lecturer It Palermo In
X74: and Vicar-Generil md prm incinl Iuquhitor in 1755. Hu Ernbryologiu Sam was
1 bent-ulltr (see Blblingrlphy) and was even translated into modern Greek for the
benefit of those Orthodox who desired to rtudy the lengths to which Reason ma the
 
Latin mind could go. Cangial-nfla'n main ion-nula was mndlnonal; Sr tn 5: upon‘.
 
ega te bapuza (even to the amnion). _HE nnxaety to bmpuse the embryo led km: to the
 
-uél-«vars; 1/2(warmer.C,1As'Ln1a-.ses'ez:za‘azrb.oz\thAb.2xo.gnud.tbedud.I\1nmntunes
 
urlier theology had been more modest, _|.nd the Rarnan Rmmb of us. contented ml:
with an edlfyzng De Betltdxrtlanefotttu xn mm malfll’. Nevertheless the eondemnanon
at‘ the unbaptlscd infant to em-na_l torment goes back to 1 \''er)_ early stage of Lima
theology and I! deeply embedded in the teazhlng of Councils. t_unta Ind popes. I! 111!
been learrledly shown by Coulton in has Rudy of Infant Perdition. The doetnne does
 
hllmnrf edi .
yméiltiamghuia this mhiect an m gum in Cohen’: boot on the Tdmud (9. m).
 
B tine‘ th ofstur. , _ _
ygunnmtignfilfdgflflmmdmgme ethic: offoeticrdelmy be hand In Armdr; Glmn:
and Hughes.
2°4
PLATE XVIII
 
.3»? .2325 3. 22$ Ea. am. :5 an» 52.3 :e_ufl§.~ 5: Ha .5.:E===. {..=:..$~ 5
 
=99!
EMBRYOLOGY IN THE EIGHT]-ZENTH CENTURY
 
 
Fig. 23. Dalerrpaliux’ dlamngr of Imman !p(nnaI0:oa (from Lemme-nimzk).
 
monsters are also very odd. But theological embryology probably
reached its climax in the report of the doctors of divinity at the Sor-
bonne on March 30, 1733, in which intm-uterine baptism by means of a
syringe was solemnly recommended. This is included.'in Deventer’s
book,‘ and has been referred to by Sterne and Spencer. For other aspects
of these tracts of thought, see Nicholle and his anonymous antagonist.
But Cangiamila and his t:olleagues—-Gerike, Kaltschmied, ete.—are
only of decorative importance to our present theme, and for fuller
information regarding them, reference must be made to the treatise of
Witkowski.' It is interesting to note that as late as 1913, 182 days was
fixed as "perfection-time," whatever that may be, by Moriani.
 
6. Ovism and Anirnalculism
 
\Ve must now return to the beginning of the century in order to pick
up the thread of the main trend of thought. By 1720 the theory of
preformation was thoroughly established, not only on the erroneous
grounds put forward by Malpighi and Swamruerdam, but on the experi-
ments of Andry, Dalenpatius’ and Gautier, who all asserted that they
had seen exceedingly minute forms of men, with arms, heads and legs
complete, inside the spermatozoa under the microscope.‘ Gautier went
so far as to say that he had seen a microscopic horse in the semen of a
 
‘ Hlsloirt dc: Aezouehmmlr, pp. 13; E
_ ' Cf._ the drscuman an Cleopatra (p. 65). Itis of interest that prenatal ba tism, even
including the use of syringes, to this day forms pm at ofiiernl Lucia theo on 1! any
nte in the Chunk of France (Ortolan). In literature, we catch tn echo of that am-
tmvemcs in the essay of Donne (I531), "That virginity is : Yerruc."
 
' Qdmy:txm' dnvnngs. r roduecd from Leeuwenhoek in Fig. 23, were nlmost
certainly : hoax; sec 1-‘. 1. Co e, PD. 63 t1’.
 
‘ Hamocknh drawing. illustrated in Fig. :4. represented not what he had um
himself but what he supposed xpemulazc: would look like if they could be u.-tn
Iu.fiiu‘:nLly clearly.
 
205
A HISTORY OP EMBRYDLOGY
 
horse (he gave a plate of it) and a similar animalcule with very large
ears in the semen of a donkey; finally, he described minute cocks in the
semen of a cock. l-laller remarks gently that he has searched for these
phenomena in vain. Vallisneri asserted the same kind of thing about the
mammalian ovum, though he admitted that, in spite of long searching,
he had never seen one. Besides the main distinction between proforma-
tionists and cpigenesists, then, there arose a division among the former
group, so that the ovists regarded all embryos as being produced from
smaller embryos in the unfertiliscd eggs, while the animalculists re-
garded all embryos as being produced from the
smaller embryos provided by the male in his
spermatozoa. The animalculists thus afforded a
singular example of a return to the ancient theory
mentioned by Aeschylus int-he 076154 (5¢€}3- 43}
Their usual view was that of Hartsoekefl and
Andry, who pictured each egg as being arranged
like the Cavorite sphere in which H. G. \l/ellsl‘:
lorers made their way to the moon, i.e. wit
:1: nap-door. The spermatozoa, like sobmany
 
' e men, all tried to occu an egg, '1‘ 13
tnlilenelggs were far fewer than splzxmtowa. 0166
were, when all was over. only 3 few MPPY
animalculcs which had been lucky enough to find
empty cggg, dimb in and lock the door behind
thegiher followers of I.ee!:x,t‘t;ni1l0¢k'f§Sb5€:'llx¢d 11141‘
there were apermatie an‘ cu es 0 o setcf.
 
4 as one oauld see by a slight difference near their
 
firlgie-121:; 0!/Inl:";,Iilkr::1: tails, that they copulated, that the females beanie
=1m"'"""’°"- rcgnant and gave birth to little ammaleu_l5.
that young and feeble ones could be seen,‘ that 31,61)’ j1¢dH':1£“' ‘hi:
and finally that some had been Observed With '_W’-if '3 5; _ :‘;;z‘1:°_
made good use, on thkes \vholc1:‘,! of lusc:::x;g,y=m D 5°°P"°‘5m'
terised all these remar 35 "0 Y °°“.l ' . -
 
The whole controversy was intimately bound up the hi:
spontaneous generation. 501’. “'1m3V°’ am “*5” ":3 . cdemmm’ mg! of
animals, if it were true that the lower ones coulh fflim at least must
slime, mud or meat infusionnfor Instance, then f :7 P for it mu“
have been made by epizmws 3”’ *3" "3 “Y °' " ‘:tar)iicture oftlut
hardly be held that a homogeneous .lnfl.1Sl0‘l"l had afiguls ‘hm thflhin
k'md_ And if gpigenesis could occur in the lower am 1
I See p. :75 -b°V=«
 
 
I Ermy, Ieet. 58.
206
EMBRYOLOGY IN THE EIGHTEENTH CENTURY
 
end of the wedge had been driven in, and it might occur among the
higher ones as well. It was in this “ray that the spontaneous generation
controversy came to have a peculiar importance for embryology in the
eighteenth century.
 
Driesch‘ has essayed to make the generalisation that all the supporters
of epigenesis were vitalist in their tendencies, while those who adhered
to the preformation theory were not. But there are too many exceptions
to this rule to make it helpful. In so far as it applies, the association
doubtless arose from the fact that the continual production in epigenesis
of new organs and new relationships between organs already formed
seemed to require an immanent formative force of some kind, such as
the vi: esrentialir of Wolff, while on the preformation hypothesis, where
ernbryogeny was little more than a swelling up of parts already there,
it could be explained simply as nutrition. But the failure of the “short-
cut" mechanistic philosophers such as Gassendi and Descartes led to
preformationism just as much as to epigencsis. A remark of Cheyne's
throws some light on this question, for in x71 5 he wrote, rallfing to
Gassendi's line of thought for different reasons, “If animals and vege-
tables cannot be produced from matter and motion (and I have clearly
proved that they cannot), they must of necessity have existed from all
eternity."’ Preformationism thus became the only resource if the uni-
versal validity of the mechanical theory of the world was to be retained.
Stahl, and later Wolff, saw no point in retaining it, and carefully joined
together what Descartes had with equal care put asunder.
 
Von Haller affords some interesting evidence against the identification
of epigenesis with vitalism and preformation with mechanism, for he
says, “Various authors have taught that the parts of the human body are
formed by a mechanism depending on general laws" (i.e. laws not
simply of biological validity), "or by the virtue of some ferment, or
by heat and cold making crusts out of the different juices, or in other
ways. All these [mechanical] systems have some resemblance to that of
M. Wolff.” Haller also always speaks of Wolff's vi: erxznlialir as "blind."
 
The original discoveries of de Grant’ and Stensen were extended by
Tauvry in 1690 to the tortoise, and by Lorenzini' in 1678 to the Tar-
pedo: so that the eighteenth century began with an excellent basis for
ovistic prefarmationism. The greatest names associated with this school
were Suarnmerdam, Malpighi, Bonnet, Von Haller, VVinslow, Vallis-
 
neri,‘ Ruysch and Spallanzani.4 But there were many others, some
of whom did valuable work, such as Bianchi,l Bourguet, Bussiere,
 
I And Ililfltiewiczz see pp. 2:4 and H9 heruftzr.
'Pfn7oxaplu':al I’h‘m-r'p!zr_ ch. a, sect. _m E. p. 6!.
' See p. 167. ‘ Ste Franchiru. ‘ D: Nat. Gem, pp. 417 H’.
 
207
A HISTORY OF EMBRYOLOGY
 
C°5°h“”‘zr Fm‘: P¢“'3\1l‘. 5i€m’-y Teichmeyer, Veroelloni, Wdussi and
Weygand: The treatises of Imbert and Plonquct were written from :51;
point ofvrew, as was the bright little dialogue ofde Houpeville. J. B du
Halftttl asserted that he could see the chick embryo in the egg beioxe
gegglisation, and Jacobaeus made a like affirmation in the case of the
 
On the other side, that of animalculistic prefor-mationism, the con-
testants were fewer. Their greatest names were Leeuwenhoek, Han.
5°€k€1’. Leibniz and the cardinal dc Polignac.’ In England the physi-
cians Keil and Cheyne supported this position, in France Geofroi and
the obstetrician la Motte, in Germany Withof and Ludwig, and in
Belgium Lieutaud. De Supenille wrote in favour of it in the Philom-
phieal Transaction: of the Royal Society, and an anonymous Swedish
work of some fame supported it. To the argument of Vallisneri that the
existence of so many aniznaleules must be an illusion, since Nature
could hardly be so prodigal, the animalculists retortetl by insraneing
such observations as that of Baster, who had taken the trouble to count
the eggs of a crab and had found that they amounted to 12,444. James
Cooke later elaborated a theory! of :1 “orld of the unborn to which the
spermatozoa could retire between each attempt to find a uterus in which
they could develop-—this avoided V:rllisneri’s argument.
 
All those other attending Anirnalcula, except that single one that is then
conceived, evaporate army, and return back into the Atmosphere again,
whence it is very likely they immediately proceeded; into the open Air, I say,
the common Receptacle of all such disengaged minute sublunary bodies; and
do there circulate about with other Sernina, where, perhaps, they do not
absolutely die, but live a latent life, in an insensible or dormant state, like,
Swallows in Winter, lying quite still like a stopped Watch when let down,
till they are received afresh into some other Male body of the proper kind, to
 
‘ Ch. 3, p. 8. _ ‘Anti-Luneriux. net. 8.
 
‘ Support: by flrolluton. Actually : urmlar argument had been E\'Dl\:d some
seventy yenn previously by F. M. van Hzln-mot, who was worrrine. not, like Cooke.
about the losr spermatozoa. but about the ovulared egg: which (nled to get fernhsrd.
Xn his strong: discourse on mttunpsychosin, he toy: (p. 15;): “Quation no. 33. More-
over. when we find that Children m the womb be formed out of Eggs. of wh-ch there
are so great . number in every woman, am we do not find one am bur: w many
Children as Ill! hath Eggs, which oh: brought into the V\'orld with her: l\lu1t we not
therefore conclude, that the rest of these Eggs were created in win. in use they should
not it some time or other main to their full perfection? Now to retrieve this dt-fficulfv.
musrwe not conclude. that the Life of these Egg: doth prvplgate xtsgli,-nothzr wry.
to the rnd that what doth not arrive or pexfecuon one tune. rmry xturn it or mother]
And um mmtm the remaining Egg: must necessarily be revolved In order to am:
perfecuon. II which in the production of than. Nature had directed her xntentrdorzi
 
In like mumer what an we euppose to be the rmon of that gxprtss commln ,
That no basnrd should enter into the ccn8f98'|"°“
 
God. which we read, Dent. 23, 2,
by means of ten llemlunons. the
 
of the Lord to the math generation; but this. that
evil might be wrought our?"
2o8
EMBRYOLOGY IN THE IZIGIITEENTI-I CENTURY
 
be again set on Motion, and ejected again in Coition as before, to run a fresh
chance for a lucky Conception; for it is very hard_to conceive that Nature is
so idly luxurious of Seeds thus only to destroy them, and to make Myriad: of
them subservient to but a single one.
 
But Cooke's attractive hypothesis, published in 1762, came too late,
as Punnett says, to save the animalculists.
 
The idea that human seed, or spermatozoa, floated everywhere in the
air (obviously derived by Cooke from the Stoic—Talrnudic-Kabbalistic
line of thought described on pp. 66, 79) led to the amusing satire of Sir
John Hill in x75o mockingly addressed to the Royal Society, Ludrm sine
Concubilu. He affected to have invented a machine for trapping the
seminal animalcules home on the ‘Nest wind.
 
Accordingly after much Exercise of my Invention, I contrived a wonderful
cylindrical, caloptrical, rotunda-concavo—convex Machine (whereof a very
exact Print will speedily he published for the Satisfaction of the Curious,
designed by Mr H-y-n, and engraved by Mr V-rtu), which, being hermeti-
cally sealed at one End, and electrified according to the nicest Lavis of Elec-
tricity, I erected in a convenient Attitude to the West, as a kind of Trap to
intercept the floating Animalculae in that prolific quarter of the Heavens.
The Event answered my Expectation; and when I had caught a suflicient
number of these small original unexpanded Minims of Existence, I spread
them out carefully like Silk~worm's Eggs upon White-paper, and then apply-
ing my best Microscope, plainly discerned them to be little Men and Women,
exact in all their Lineaments and Limbs, and ready to ofier themselves little
Candidates for Life, whenever they should happen to be imbibed with Air or
Nutrimcnt, and conveyed down into the Vessels of Generation.‘
 
On the experimental side, Garden came forward with descriptions of
little men inside the animzilcules, thus “confirming" the work of Gautier
and Hartsoeker. It is fair to add, however, that Garden held quite
enlightened views of the mutual necessity of egg and sperrnatozoon. So
did Massuet, whose dissertation appeared in 1729 at Leiden. An
animalculist, he yet believed both egg and sperm to be needed in
generation, the fomier rather as a nidus. He reversed the Malpighian
view. Si ovum gallium, he said, non foenmdatum minorrnpio inspexrfis,
mzlla in ca animalirfanna apparebit. He gave the correct explanation of
Nuck’s experiment (see p. 163), saying that :emx'm'.r aum seu animaltula
had first passed up the Fallopian tube. In his plate he figured spemta-
tozoa, chick embryos and tadpoles all together, confusing the former
with the primitive streak.
 
Another adherent of enlightened common sense “as Hugh Cham-
berlen, one of the famous obstetrical family. In his English translation
 
1 Se: 1!. 4:.
u.l.—u 209
A HISTORY or EMBRYOLOGY
 
(1683) of Maux_iceau‘s midwifery (was) he took exception to th.
doubIe—seed (Epicurean) opinions of the French writer, and addcd the;
following as a footnote:
 
. Our author lying under a Mistake, in his notions concerning the Testida
in this chapter, I shall here give my sentiments. We find that the Tcsticles
of a Wornanare no more than, as it were, two Clusters of Eggs, which [ie
there to be impregnated by the Spirizuous Particles, or Animating Emu.
VN1115. Oonveyed out of the Womb through the two Tubes, called by our
Author, Defcrent Vessels. . . . Some days after the impregnation of the Egg
or Eggs, as in Twins, they decid through those two Tubes into the Womb:
where being placed, the Embrio takes up its quarters.
 
But la l\Iotte maintained that the egg (which he identified with
the Graafian follicle) was too big to go down the Fallopian tube,
launxrrll‘ Sbaragli, another writer on the animalculist side, agreed with
As for the supporters of cpigenesis, they were few, but they included
Desmrtes, de Maupertuis, Antoine Maitre-Jan and John Turberville
Needham. 1‘/Iinor Writers on the same side were Tauvry, lVelsch,
Dartiguelongue, Béttger, Drelinourtius and Mazin. After r75o C. F,
Wolff brought an abiding victory to their opinion.
 
Among the arguments brought forward against the preformationists
‘VETS 3
(I) That it is impossible to explain the production of monsters on a
preformation theory. Brunner first brought this point forward in 1683.
but its classical statement was that of Etienne Geolfroy de St Hilaire, in
his work of 1826 on experimental teratology. On its history, see Strohl
and Dareste.
 
(2) That preformation is incompatible with the facts of regeneration.
An intelligence, argued Hartsoelser in 1722, that am reproduce the lost
claw of a crayfish, can reproduce the entire animal. This point much
impressed Erasmus Darwin.‘ The whole subject was much to the fore in
the eighteenth century owing to the brilliant observations of de
Réaumur and Trembley!
 
(3) That the extraordinary resemblance between small embryos of
mammals, birds, reptiles, etc., discredits preformation. This was the
View of Prévost 8: Dumas, 1834-1838.
 
Some maintained a quite independent position, such.as Bufion, who
welded together an epigenetic theory of fertilisation with a prtfnrmr
tionist theory of embryogeny. Pascal (not the great Jansemst) W‘ ‘W’
'Zoonoam'n, :, 45:137. ' See Mer-
 
zro
EMBRYOLOGY IN THE EIGHTEENTH CENTURY
 
ward dte chemical view that fertilisation consisted in a combination
between the acid semen of the male and the “lixivious” semen of the
female, perhaps because in chemistry acids had been regarded as male
and alkalis female.‘ Claude Perrault‘ and Connor also suggested that
the formation of the embryo was a fermentation set up in the egg by the
sperxnatic animalcule. In this they were following the example of van
Helmont, who had originally suggested such a theory.’ In 1763 Jacobi
discovered how to fertilise fish eggs with milt; a practical matter which
had a good deal of influence on biological theory. De Launay alone still
held to the Aristotelian conception of form and matter.
 
7. Spontaneous Generation
 
There is no need here to do more than glance at the spontaneous
generation controversy itself, for it has long been well known in the
history of biology, especially in connection with the subsequent work
of Pasteur. J. T. Needham’s books, New I|rIicra.tco1>1‘:ul Dircoverie: of
1745 and Obrmvnliant upon the generation, composition, and decomposition
of Animal and Vegetable Subxtmices of 1749. exercised a considerable
influence. They were written concisely after the French fashion (Need-
ham had been educated at Douai), and with some brilliance of style,
and it is hardly true to say, as Rad] does, that their experimental
foundation was meagre. That it was inadequate was proved defi-
nitively as events tumed out by Spallanzani. De Kruif's picture
of the controversy is false and misleading, especially in its estimate
of Needham, “ho is much more truly described in the words of Louis
Pasteur.‘
 
Need.ham’s case rested upon the statement that if meat broth was
placed in a sealed vessel and heated to a high temperature so that all life
in it was destroyed, it would yet be found to be swarming some days
later with microscopical animals. All depended, therefore, upon the
sureness with which the vessel had been scaled and the efficacy of the
heat employed to kill all the animalcules initially present; and in the
 
ensuing controversy Needham lost to Spallanzani entirely on the question
‘of technique. It may be remarked here without irrelevance that the
problem is still in a sense unsolved, for what the experiments of Sp:illan-
zani proved was that animals the size of rotifers and Protozoa do not
 
"2153. Gregory‘; interuting essay on the minute and mechanical model: of
193'» s
 
' One:-m diverm de Pluyriwe t1|1!cnru'qu4(i7z ), nl. 2, . 439.
 
' Omu, l, :1. How intercatrdtthcse men vvouldIha‘\"l: bee: in modern researches on
the changes at fertilisation of respiration, glyeolytie activity, and enzyme action in the
prmoplum of the egg cell.
 
* For an accurate and detailed account at’ the controversy, see Prescott.
 
211
A IHSTORY or IZMBRYOLOGY
 
originate s ontaneousl
 
pasta“. “.3: that orgmyivs :03’ Ptl'ov_cd. by lgose of
in that way. The knowledge which has been at uirrtili yngmme v 2 now
"filter-passing organisms," such as the niosaii V"ll'|1S"i.)ir'e1ci::"t}§a:s of
Pl-'15‘. and phenomena such as the bacteriophage of Twort and d’iilei:ll:
has re—opened the whole matter, so that of the region between for
example, the semi-living particles of the bacterioph:lge(1°—i: Em)’ and
the larger-sized colloidal aggregates (mm gmm) We know ,cma,k3M
little. The possibility of the new formation of viruses without s
ancestry, in the cells of living hosts, is to-day an open qucsting aid we
still have no proof that their origin from non-living Organic ',m,c,;31
can never occur. Recently it has proved possible to "synthesise” infec-
tive viruses from separate protein and nucleic acid oonstituents of other
viruses, constituents some of which alone are incapable of reproduction.‘
The dictum onm: z-ioum 2: viva, accepted with such assurance by the
biologists of the early twentieth century, may thus tum out not go mum
quite what they thought.
 
But to dwell further on this would be a digression. The important
point was that Spallnnzani's victory was a victory not only {or those uho
dishelievcd in spontaneous generation, but also for those who believed
in the preformation theory of embryogenyf By 1786, indeed, that view-
point was so orthodox that Scnchier, in his introduction to an edition
of Spallanzani’s hook on the generation of animals and plants, could
treat the epigenesists as no better than atheists.
 
Spall-anzani's views on embryology were largely drawn from his study
of the development of the frog’s egg (see Fig. 19). Here he went far
beyond Bose, but in spite of many careful observations he thought he
saw the embryo already present in the unfertilised ovum. This led him
to claim that Amphibia ought tobe numbered amongviviparous animals.
His principal step forward was his recognition of the semen as the actual
agent in fertilisation on precise experimental grounds—the narrative of
his artificial insemination of a bitch is too famous to quote: he said it
gave him more intellectual satisfaction than any other experiment he had
ever done. This demonstration finally dbposed of the aura .mm'naIr'.t
wlvich Harvey had. found himself ohliged to adopt on the grounds of his
dissections of does. But Spallanznni failed to convince himself that the
 
spermatozoa themselves were the active agents.
 
‘ Fl-aenlrcl-Canrat has given : good description of this work. There now seems no
 
mson why it should not be possible In the {ensemble future to nynqmiie . nucleic
hid: when introduced into n host cell
 
acid from other organic izhemiaal ntlnmnes w _ _
would show infecuvity, i.:. n self-rephaxtion or reproduction within the metabolic
 
' - This was realised at the nine rspecially by Patrin, who am to Needhanfa defence
 
in 1778 (see Btlikiewiez, p. I47). See on, p. 218.
212
EMBRYDLOGY IN nu»: EIGHTIZIZNTH CENTURY
8. Preformation and Epigenesis‘
 
Of all the preformationists Charles Bonnet was the most theoretical.‘
He was an adherent of that way of thinking mainly on the theoretical
ground that the organs of the body were linked together in so intimate
a manner that it was not possible to suppose that there could ever be a
moment when one or two of them were absent from the ranks.
 
One needs [he said] no Morgagni, no Hallcr, no Albinus to see that all the
constituent parts of the body are so directly, so variously, so manifoldly,
intertwined as regards their functions, that their relationship is so tight and
so indivisible, that they must have originated all together at one and the same
time. The artery implies the vein, their operation implies the nerves, which in
their turn imply the brain and that by consequence the heart, and every
single condition a whole row of other conditions.
 
Bonnet compared epigenesis to crystal-growth, in which particles are
added to the original mass independently of the plan or scheme of the
whole, i.e. in contrast with the growth of an organism, in which particles
are added on only at certain places and certain times under the guidance
of "forces (It: rapport.” Przibram has recently discussed the question of
how far such at comparison is admissible, but in Bonnet’s time at any
rate it became very famous. Bonnet referred to von Haller’s dis-
covery of the intimate relationship between embryo and yolk as evidence
for his theory. The embryo begins, according to him, as an exceedingly
fine net on the surface of the yolk; fertilisation makes part of it beat and
this becomes the heart, which, sending blood into all the vessels, ex-
pands the net. The net or web catches the food particles in its pores, and
Bonnet supposed that if it were possible to abstract allthe food particles
at one operation from the adult animal, it would shrivel and shrink up
into the original invisible web from which it originated.
Bonnet was no more afraid of the emboflement principle than was
 
Haller; indeed, he called it “one of the greatest triumphs of rational over
 
‘ \Vhite'| 77!: Phlagittm Thzovy will interest those who Willi to study the rather
striking panllel between chemistry and biology in the eighteenth century. Broadly
speaking. rationalism in science had too much got the upper hand of zrnnincism. The
trace: of preforrnntionism remaining in modern biology are well reviewed by Huxley
8: de Beer. \Vhitrnan distinguished between "predeterrninntion,” I physiological or
potential prcfnmistion not capable of rniuoscapic resolution, and "predelmeation,"
which I! the old morphological or more preformation. Modern embryology might
therefore be called "predzterm' ed epigtnesis." “Inddington has recently (.952) intro-
duced the term “e~pigeneiies" t nelnde everything etmctrned \\llh the causal analysis
 
of development, that is to say, with the genes Ind thtir effects in embryonic life as well
as the merphogenetie rnedunisms themselves.
 
' Bonnet‘: nu.-tiul contributions to science were not numerous but rather impor-
tani. ll: Congflfled in 1745 (r779, 13. 36) Leeuwenhoek‘: dI5C<7VE1'y of the pnnhrno-
gum: or aphids (1702), and he announced the formation or new individuals alter
the cutting of worms into regrntrits (nee the paper of Erhard). The value of the former
 
observation I! 1 support for ovism is evident. For further dcuil: concerning him.Ie:
Lemaitre and Whitman.
 
2x3
A HISTORY or EMDRYOLOGY
 
sensual conviction." Many of his argum -
Haller's, and he says in his preface that heehziil i:eri:etehEhisml2::li°snosuif
time before Haller's papers on the chick appeared but then finding his
. . . .
 
own views confirmed by the expenmentally better founded ones of
Haller, he determined to publish what he had set down. Thus in one
place he says,
 
I shall be told, no doubt, that the observations on the development of the
chick in the €88. and the doe in the maternal uterus, make it appear that the
parts of an organised body are formed one after another. In the chiclr for
instance it has been observed that during the early pan of incubation the
heart seems to be outside the animal and has a very different form to nhat it
will have. But the {eehleness of this objection is easy to apprehend. Some
people wish to judge of the time when the parts of an organised body begin
to must by the time when they become visible to us. They do not reflect that
minuteness and transparency alone can make these Pang inyi5,'],1¢ gg ‘,3
 
although they really exist all the time.
 
Bonnet was therefore what might be mlled an "organic preforma-
tionist," for his objection to epigenesis lay in the {act that it did not
seem to allow for the integration of the organism as a whole. His
mistake was that he assumed the capacities of the adult organism to
be present all through foetal life, whereas the truth is that they grow
and dilferentiate in exactly the same way as the physical structure itself
does. Bonnet's philosophical position, which has been analysed by
\Vl1itm:1n, again contradicts the generalisation of Dr-iesch' that all
the epigenesists were vitalists and all the preformationists mechanists.
For Bonnet an epigcnetic and a mechanical theory wen: one and the
same; he hardly distinguished, as Rédl says, between Descartes and
Harvey; and it wasjust the neodritalist idea of the organism as A whole
that he oould not fit in with epigenesis. Needham and \Voll1' were un-
doubtedly epigenesist-vitalists, and Bonnet was undoubtedly a pre-
fonmtionist-vitalist, but Maupertuis was equally clearly an epigensisb
mechanist.
 
G. L. Leclerc, Comte de Bufion, the most independent figure in the
controversy, stood alone as much bemuse of his erroneous experiments
as because of his originality of mind. As has so often been observed.
Bufion was not really an expcrimentalist at all; he was a writer, and
preferred other people to do his experiments for him. The vol'ume.0n
generation in his Hirtoin Natxmlle begins with a very long historical
account of the work which had been done in the previous centuries on
embryology. At the beginning of the section on reproduction in general
he said,
 
I And gr Eilikiewicz.
214
EMBRYOLOGY IN THE EIGHTEENTH CENTURY
 
The first and most simple manner of reproduction is to assemble in one
body an infinite number of similar organic bodies, and to compose the sub-
stance in such a manner that every part shall contain a germ or embryo of the
same species, and which might become a whole of the same ltind with that of
which it constitutes a part.
 
Such an idea resembles the ancient atomistic speculations, and was
explained by VV. Smellie, the obstetrician, who translated Bufion i.nto
 
English, as follows:
 
The intelligent reader will perceive that this sentence, though not very
obvious, contains the principle upon which the whole theory of generation
adopted by the author is founded. It means no more than that the bodies of
animals and of vegetables are composed of an infinite number of organic
particles, perfectly similar, both in figure and substance, to the whole
animal or plant of which they are the constituent pans.
 
This conception explains l3ufl'on's curious attitude to the preformation
question. An embryo was preformed in its germ because all the parts
of the germ were each a model of the animal as a whole, but it was also
formed by epigcnesis because, the sexual organs being first formed, all
the rest arose entirely by a succession of new origins. Bufi'on’s “organic
living particles” bear some resemblance to the “biogen molecules"
which later generations were to discuss,‘ and he says that an exactly
similar but simpler stnrcture is present in dead matter.
 
In his discussion of former theories he resolutely rejects the ¢rnI7oi!e-
merit aspect of preformationism, giving various calculations to show
 
its impossibility and maintaining that
 
every hypothesis which admits an infinite progression ought to be rejected
not only as false but as destitute of every vestige of probability. As both the
vermicular and ovular systems suppose such a progression, they should be
excluded for ever from philosophy.
 
He completely destroys the theory which the ovists and animalculists
had set up in order to explain resemblance to parents, namely that
although the foetus might originate either from egg or spermatic animal-
cule originally, it was moulded into the form of its parents by the in-
flucnce of the rnatemal organism during pregnancy. This field,’ which
was more than once disturbed by the contestants during the course of
the century, received systematic attention from time to time by medical
writers. There was a memorable dispute on the point between Turner
and Blondel, whose polemics, written in an exceedingly witty manner,
are still very pleasant and amusing to read. Blondel was the soeptic and
 
‘ And even more to the Item: of Annsgoru and the Stoic-Knbbslistic "ends."
‘ See p. :9.
 
215
- A msromr or EMBRYOLOGY
 
 
Fig. 25. G. L. L. J: Bujon and hi: frialdr xtudying marnnmlian generation.
71m mm an m n 1-ignem in Buflovf: "Him-nre Nanu=l.le" at 11:: head qf
 
pr um _
5i"f.i2?£  "¥‘5e“1s‘I’2'f71”.‘~§?'.§,‘3{
 
MR7! Lwiv Ddvbwtvl (1715-I799) loglu dawn the miaaxtope. 77:: mrgm in atttnd.
one: may be miner Gutnnzu 4. Jormz-emu (1720-1785) or 2‘. F. Dubbard (I703-1779),
mo mm eolhbarman af B_uflon. /in um uwlurr mu epzgemm and anmuzl.
mhm, and mug in mm mumk: Whit]! ha: nnw bun Md, they mm am-inmx
that M luzdfmmd mmmnnzaa in the Gme/Ea: fulluks of xhef.-male ovary. The um:
xhomx urnnrining the mmmmluza generative organ: (me pp. 168 am! an: of the
"Hisroire Naturelle," Va!‘ 11, r75a). The ingmxhur xdznofizarwn of the pawn is due
to Pmfeuar R. C. Pu-rmenr on Guéneau dz Alonlblliard, su Bnmel and Mmquae
 
Turner the defender of the numerous extraordinary storiu which
passed for evidence on this subject. It is interesting to note that Turner
believed in the continuity of foetal and maternal blood-vessels. Krause
and Ens later supported the opinions of Turner, while Okcs, in 3 Cam-
bridge disputalion, argued against them.‘
 
BuEon's sixth chapter. in which he relates the progress of his on
experiments, is unfortunate, in that his main result “as to discover
spermatozoa in the liquor fulliculi of ovaries of female animals (see
Fig. 25). The explanation of how he came to make such an egregious
mistake has never been satisfacton'Iy given, and it was not long before
the truth of the observation was questioned by Lederrnuller} It led him
naturally to the assertion that the ovaries of mammalia were not cgg~
producing organs but animalcule—producing orgam, and to the Via!’
 
I ' kn tha h hl h H lorrunittedhizrulfmmme
.,..i§i‘Z§i§‘.f.‘.".i.’3‘..l'.".,.. i’§Z"--p.,‘$..-S .’?...’$-“Er Zimfi inarmx mm; (Plain-
 
mphy oflllfnd, pp. 235.).
' Vcmath. p. 13.
216
EMBRYOLOGY IN THE EIGHTEENTH CENTURY
 
that the beginning of embryonic development lay in the fusion of the
male with the female spermatic anirnalcules——a curious revival of Epi-
cureanism.‘ But it is to be observed that he does not mean one male
animalcule with one female animalmle, but rather all with all, in a kind
of pangenesis.
 
All the organic particles which were detached from the head of the animal
will arrange themselves in a similar order in the head of the foetus. Those
which proceeded from the backbone will dispose themselves in an order
corresponding to the structure and position of the vertebrae.
 
And so on for all the organs. The fact that for the organs common to
both sexes a double set of nnimalcules will thus be provided does not
give Button any difiiculty and is fully admitted by him. Accordingly he
could only agree to the aphorism «mm: vivum ex aw in the sense of
Harvey namely as referring to the egg-shaped chorion of Vivipara, and
definitely not in the sense of von Baer, namely in the modern sense.
 
Eggs, instead of being common to all females, are only instruments employed
by Nature for supplying the place of uteri in those animals which are de-
prived of this organ. Instead of being active and essential to the first im-
pregnation, eggs are only passive and accidental parts destined for the
nourishment of the foetus already formed in a particular part of this matrix
by the mixture of the male and female semen.
 
Biology at this period was still labouring in the dark without the
illumination of the cell-theory, and therefore unable to distinguish
between an egg and an egg-cell.‘
 
In spite of his leanings towards epigenesis, Bufion repeated precisely
the error of Malpighi.
 
I formerly detected [he says] the errors of those who maintained that the
heart or the blood were firs: formed. The whole is formed at the same time.
We learn from actual observation that the chicken exists in the egg before
incubation. The head, the backbone, and even the appendages which form
the placenta are all distinguishable. I have opened a great number of eggs
both before and after incubation and I am convinced from the evidence of
my own eyes that the whole chicken exist: in the middle of the cimtrix the
moment the egg issues from the body of the hen. The heat communicated
 
‘ it is interesting that the rejection of the Epicurean theory of female seed by the
Latin theologians led, in 2ighteenth—cenl'ury moral theology, to 3 very unequal em-
phasis on mumrbuion u I am in main and females. The male (cf. the influence of
the nnimnlcuhsts) was to be regarded as little short of I murderer if tfluxio Irwin’!
occurred; the female could not thus -in quiz: rennn semenin mxlmfbur nan datunuee
CI‘pellmn.nn. pp. 88 R‘.
 
In 1778 \V. Cruikshank found blasxoeystl in Ll’): Fallopian tubes of the nbbit on
lhe third day utter coitus, but his -neunte ohm-vuim-is were not publnhed till 1797.
 
217
A ms-roar or EMBRYOLOGY
 
2:5‘ 5): inculgltian czrpaods the pam only. But we have never been able to
°"“"" ‘“ “"-W“? what parts of the foetus are Erst fixed, at the mo.
 
ment of its formation.
 
The expenmcnt oftaking a look at the cimtrioes of eggs on their way
down the parental oviduct seems so obvious that Bulfon may well have
thought of it, and it would he really interesting to know what factor
in the intellectual climate made him regard such an observation as
not worth attempting. His obsenations on the embryo itself were good
and in some “my: new; thus he noticed that the blood firs: appears on
the "plaoenta” or blastoderm, and for the first few days seems hardly to
enter the body of the embryo. He gave an extremely good account of the
whole developmental process in the chick and in man, and his opinions
on the use of the amniotic liquid and the functions of the umbilical
oord were advanced.
 
J. T. Ncedham, however, spoke very clearly in favour of epigenesis,
though he himself did no emhryological experiments.‘ His Idéerommairz
of 1776, written against Voltaire. who had called him a Jesuit and
who had drawn materialistic inferences from his writings, contain the
 
following passage:
 
The numerous absurdities which exist in the opinion of pre—existent germs,
together with the impossibility of explaining on that ground the birth of
monsters and hybrids, made me anbrace the ancient system of epigenesis,
which is that of Aristotle, Hippocratns, and all the ancient philosophers, as
well as of Bacon and a great number of savants among the neoteriqucs. My
 
obsen-ations also led me directly to the same result.
 
Needham’s embryology is mostly contained in his Obrmwtimu nou-
celler run In Griréralinn of 1750. He was explicitly a Leibnizian and
 
postulated a vegetative force in every rnonad.
 
Needham was not the only thomughgoing epigenesist of this period.
Maupertuis, whose Vina: P}1_)'s1'que was published anonymously in 1746,
mme out very clearly in {armor of this doctrine.‘ He wrote:
 
I know too well the faults of all the systems which I have been desc-n'bing
to adopt any one of them, and I find too much obscurity in the whole matter
to wish to form one of my own. I have but a few vague thought: which I
propose rather as thoughts to be examined than as opinions to  received,
and I shall neither be surprised nor think myself aggrieved if they are
 
I Hi: valuahle biologial discoveries have beat samewim mnhadawed by th«
lpontaneaus geuention controversy. In fact he made ldrnirabée mnueigptrai-slang}:
cu-nplzmted physiology, sexual and gains], ofcephalapods -n -.-1].;-11> 5;, mum in
pollen grains is maloguu oiepenmtozoa and was the lingo nee mvm
 
them, and he described the horned eggs of damiobraneh bu.
 
- Se: Brunet.
218
EMBRYOLOGY IN THE EIGHTEENTH CENTURY
 
rejected. It seems to me that the system of eggs and that of spermatic animal-
cules are both incompatible with the manner in which Harvey actually
saw the embryo to be formed. And one or the other of these systems seems
to me still more surely destroyed by the resemblance of the child, now to the
father and now to the mother, and by hybrid animals which are born from
two different species. . . . In this obscurity in which we find ourselves on the
manner in which the {oetus is formed from the mixture of two liquors. We
find certain facts which are perhaps a better analogy than what happens in
the brain. When one mixes silver and spirits of nitre with mercury and water.
the particles of these substances come together themselves to form a vegeta«
tion so like a tree that it has been impossible to refuse it the name.
 
This was the Arbor Dianne, which played a great part in the embryo-
logical controversies of the eighteenth century. It has much interest
for us, for it was perhaps the first occasion on which a non-living
phenomenon had been appealed to as an illustration of what went on in
the living body. It is true that Descartes long before had said that the
movements of the living body‘ were carried out by mechanisms like
clocks or watches, and that they resembled the statues in certain gardens
which could be made to perform unexpected functions by the pressure
of a rnanipulator’s foot on a pedal, but these instances were all artificially
constructed mechanical devices, whereas the Arbor Dianae was a
natural phenomenon quite unexplained by the chemists of the time,
and the lineal forerunner of Lillie’s artificial nerve, and Rhumbler’s
drop of chloroform. VVe know now that its formation is a simpler
process than anything which occurs in the developing embryo, but the
growth of knowledge has made it undeniably clear‘ that the same forces
which operate in the formation of the Arbor Dianne are at work also in
the developing embryo. To this extent Mztupertuis is abundantly
justified, and Driesch’s comments on him are not in agreement with the
facts. Maupertuis continues:
 
Douhtless many other productions of a like kind will he found if they are
looked for, or perhaps if they are looked for less. And although they seem to be
less organised than the body of most animals, may they not depend on the
same mechanics and on similar laws? Will the ordinary laws of motion
sufiice, or must we have recourse to new forces? These forces, incomprehen-
sible as they are, appear to have penetrated even into the Academy of Science:
 
at Paris, that institution where so many opinions are weighed and so few
admitted.
 
Maupertuis goes on to speak of the contemporary deliberations on the
subject of attraction.
 
* See (2.; zxlmyle Rinne or Puibnrm
2 I9
A HISTORY OF EMBRYOLOGY
 
Chymistiy has felt the necessity of adopting this conception and attraaire
force is nowadays admitted by the most famous chyrnists who have carried
the use of it far beyond the point which the astronomers had reached. lfthis
force exists in nature, why should it not take part in the formation ofaniimlsi
 
Maupertuis was thus an epigenesist and a mechanist at the same time.
His opinions have an extremely modern ring, and his only retrograde
step was in suggesting that the spermatic animals had nothing else to do
except to mix the two seeds by swimming about in them. But that legacy
of ovism was common all through the eighteenth century, and thirty
years later Alexander Hamilton could still say, “From the discovery of
Anim.-ilcula in ::mine man-i:li'rio by Leeunenhoclfls Glasses, a new
Theory was adopted which is not yet entirely exploded.”*_ _
 
But the real middle point and fulcrum of the whole period lay in the
controversy between von Haller and Caspar Friedrich Wolff, the former
at Gottingen and the latter at St Petersburg in the academy of Queen
Catherine. Kirchhoff has described this polemic. \Volfi‘s Theorm
Generntioni}, uhich was :1 defence ofepigenesis on theoretiml and philo-
sophical grounds, uritten in a very forrml. l9g‘°-‘ll and ‘"‘_’“‘d“bl°
manner, appeared when he was only tW¢Df}"5'-‘ 3"-'3" Pld» "1 *759-
Leilmiz, as Ridl points out, had borrowed from the earher preforrna-
fionisrs the conception of a unit increasing in bulk in Order *0 590911“
another kind of unit, but W013. f01_l°W!_"8 N¢‘dh“m3 b°"°‘"d rm";
Leibniz the idea of a inonad destlopmg into an urg=1“‘5‘.“ 5}’ 335"“;
its own inherent force, and to this he joined the Stahlialn 31011:; ililra
supra-physical generative force in nature. On the Pl’-‘Cm’ V5‘ 5- °_ I 3
work was indeed of the highest lmP°mn°°‘ If the embfio mtvs 5'
he argued, if all the organs are actually_prcsent 3! ‘I13 V") ¢3'_1°5‘ 5'38!‘
and only invisible to us even with the highest power; of our m}icr0S0l:P€I3]:
then we ought to see them fully formeitl as soon as we see I em a ha“;
in other words, at the moment a1 “'l“‘h ‘"7 gm“ om: Ecmcjf the
view, it ought to have the form and shape. though “_°‘l; 35”“: um“
same organ when fully completed in the embryo at hirt n ‘hm cm
band, if this is not the way In which du'c1cpm=n‘hn8°°‘-‘S; . into
ought to be able to see uith the microscope anti 5 Pceh onegggemm
another shape. and in fact a series of ap_pe:u'.n_iCcS» :1 words 2 Suit:
{mm um which imimdmely pmcfides 1‘, or utitlile “riinitive emhr)“
of advancin§hfi“l;'pt:lllJ::sa:fllil-gtfivl'::lT;lp;|rst: he bloliid-vessels of (ht
oriic rn:isS- 0 C - .
bkgstodgfin in the chick. for he saw that at one _XflC|:1l'zi1‘:nx:()lth;JSe€clI]:-fl1}l’I?lls
‘"5 “'35 in "~‘55“’"°°’ Wm‘ die mama“ baht’: "that the homogene-
mgcmsggpicaheseaxches led turn to the conclusion
 
I Ekvnfflli, P. 43.
220
EMBRYOLOGY IN THE EIGHTEENTH CENTURY
 
ous surface of the blastoderm partially liquefies and transforms itself at
these points into a mass of islands of solid matter, separated by empty
spaces filled with a colourless liquid but afterwards with a red liquid, the
blood. Finally, these spaces are covered with membranes and become
vessels. Consequently it was demonstrable that the vessels had not been
previously formed, but had arisen by epigenesis.
 
Holler replied to this new experimental foundation of morphogenesis
without delay, for he was working on the development of the chick at
the same time, and held closely to the opposite theory. \Ve have already
seen what his one and only argument against \Volff was. He used it time
after time in all its possible variations, maintaining stoutly that the chick
embryo was so fluid in the early stages that \Volif had no right to deny
the presence of a given structure simply because he could not see it.
Haller’s explanation of Wolff’s results was that the blood-vessels had
been there all the time but that they had not become visible until the
moment at which Wolff saw the islands forming.
 
After I had written the above [said Hallei-], M. Wolff made new objections
against the demonstration. Instructed by new researches, he denies absolutely
that the yolk-membranes, which he makes two in number, exist before
incubation. He pretends that they are new and that they are born at the be-
ginning of incubation, and consequently that the continuity of their vessels
with the embryo does not in the least prove that in the body of the mother
the yolk received vessels from the foetus. I have compared the observations
of this great man with my own and I have found that the yolk never has more
than one pulpy and soft membrane, part of which is called the umbilical
area, and that the fine exterior membrane does not belong to the yolk but to
the inner part of the umbilical membrane. . . . I do not believe that any new
vessels arise at all, but that the blood which enters them make them more
obvious because of the colour which it gives them, and so by the augmenta-
tion of their volume they become longer.
 
Wolff replied by another extensive piece of work, which he called De
Formation: Intertinorum, and which appeared in one of the publications
of the Russian academy for 1768. It mined preformationism. In it he
demonstrated that the intestine is formed in the chick by the folding
back of a sheet of tissue which is detached from the ventral surface of
the embryo, and that the folds produce a gutter which in course of time
transforms itself into a closed tube. The intestine, therefore, could not
possibly be said to be preformed. From this as a starting-point Wolff
went on to propose an epigenetic theory which applied the same process
to all organs. It is interesting to note that the facts brought forward
 
by Wolff have never been contradicted, but have been used as a founda-
tion to which numberlcss morphological embryologists have added
 
22!
A nrsronv or EMBRYOLOGY
 
facts discovered by themselves. It is noteworthy too that although
Wolff 5 second general principle, that of increasing solidification during
embryonic development, led to no immediate results, it has been
abundantly confirmed since then. His observations on the derivation of
the parts of the early embryo from "leaf-like" layers were even more
important, and acted as a very potent influence in the work of Pander
and von Baer.
 
It happened, however, dint Haller had much the greater influence in
the biological world at the time, so that Wold": conceptions did not
immediately yield fruit in any general advance. Looking back over the
second half of the seventeenth and the first two-thirds of the eighteenth
centuries, it is remarkable how little theoretical progress was made in
View of the abundance of new facts which were discovered. Punnett,
in an interesting paper, has vividly brought this our.
 
The controversy between the Ovists and Artimalculkts had lasted just I
oentury, and it is not uninteresting to reflect that the general attitude or’
science towards the problem of genetation was in 1775 much what it had been
in 1675. When the period opened, almost all students ofhiology and medicine
were Prefomiationists and Ovisrs; at its close they were for the most part
Ovists and Prefotznationists.
 
Ovisrn sprang in the first instance from de Gra.1l"s discovery of the
mammalian “egg,” which gave a new and precise meaning to Harvey's
aphoristn. Preformationism, already old as a theory, acquired an
apparent factual basis in the work of Malpighi and Swammerdam, and
allied itself naturally with twism. With Leeuwenhoek and his sperma-
tozoa, animalculism txrne upon the field. The main outlines ofthe battle
which went on between the two viewpoints have already been drawn,
but it is worth remembering that there were independent minds who
were impressed by the obvious facts of heredity and found it difficnlt to
call one sex essential rather than the other. Among these Needham and
Maupertuis‘ may be counted. Among the lesser writers, James Handley
with his Jlletlmnical Ermyr on the Animal Oemrwmy of 1730 ought
to receive a mention. Though fond of theological arguments, ‘he
upheld the common-sense attitude against ovists and anirmlcullsfs
a‘.ike——“\Ve dissent in some things," he said. "bath from Lceuwcnhoeck
and Harvey. . . . Both the semen and ova (notwithstanding all “His 1‘-‘in
be said) we believe to he a mum sine qua mm in every Generation. But
what finally killed anirnalculism was the discovery in so many’ Pl-‘"5 °i
small motile living beings. flagellates, Protozoa. l-"E5 V‘l7"°3' 1‘ ‘V13
difiicult to maintain in the face of this new evidence that the sperma-
I See the excellent review or Glass which describes. inler olia, zuruv=fl“"' xfivnm
investigation of polydanyly in mm.
222
EMBRYOLOGY IN THE EIGHTEENTH CENTURY
 
tozoa were essential elements in generation, though the seminal fluid
itself might very well he, as of course was Spallanzanfs opinion. The
prefomiatian theory was what was holding up further progress, and
when VVolff’s arguments prevailed in the very last years of the eight-
eenth century, the way was open for the recognition of the true value
of the spermatozoa.
 
The physician rl'Aumont, otherwise unknown, who wrote the article
on "Generation" in Diderot‘s famous Emyrlopaediu, brought this out
in an interesting Way; for himself an ovist he summarised the arguments
which in 1747 were destroying the anirnalculist position, and reducing
rapidly the number of its adherents.
 
1. Nature would never be so prolific as to produce such millions of
 
spermatic animalcules, each one with its soul, unnecessarily.
 
2. The sperrnatic animalcules of all animals are the same size, no
 
matter how large the animal is; how therefore can they be in-
 
volved in its generation?
3. They are never found in the uterus after coitus, but only in the
 
sperm.’
4. How do they reproduce their kind? -
5. \Vhat evidence is there that they are any different from the
 
animalcules (of similar shape, etc.) which are to be found in hay
infusion, scrapings from the teeth, and many other places? No one
supposes that these have relation to reproduction (Bourg'uet).’
 
9. The Closing Years
 
The last forty years of the century were not marked by any great
movement in a fruitful direction for morphological embryology, an
iconographic wave of some merit due to Albinus, W. Hunter, Tarin,
Senfi, Rosenrnuller, Danz and Sdn1mering' excepted; and it was not
until 1812 that J. F. Meckel the younger translated Wolfl"s papers into
German. This was one of the principal influences upon Pander and von
Boer. In his introduction, Meckel describes how \Volfl"s work had been
disregarded, and points out that Oken, writing in r8oG, had apparently
never even heard of it. In the very early years of the nineteenth century
morphologiml embryology received a great impetus however. One of
the most interesting figures of the new period was de Lézérec, a Breton,
whose father had been in the Russian naval service. The son, as a Russian
 
_ ' Loss of motility, and agglutination, doubtless disguised their presence from the
investigators of this riod.
 
' \’nn Baer hirns¢lr(I8z7) believed in the extraneous nature of the xnimaleules, and
attempted to zxprtss it by the name “Ape:-rmtoeoa" which he gave them. Not until
Kfillsilterg work in r8.u wu theirhistogermu as non-ml |issue—¢l:mmn dernonscmed.
 
ee st.
 
223
A HISTORY or EMIIRYOLOGY
 
ntaxéal ;adet,b no doubt stunulated by the writings of .\vo1t-2, who had piml
ah ‘ 5"“ _‘-"E. usedlo incubate eggson board slup. He eventually left
t  sea, studied medicine at Jena, and in 1808 wrote an excellent disser-
tation on the embryology of the chick, which Stieda has recently brought
to light. He then went to Pans, and taking a medical appointment at
G“3d¢l°‘-‘PC. was lost to science. Very much more important was the
work of Pander in 1817 and von Baer in 1828,‘ but it belongs to the
modem penad,' and must be left for the next volume. Here it will
sufliee to say that these great investigators established firmly the con-
ception of the germ-layers (our now familiar ectoderm. mesoderm and
endoderm) and clearly distinguished between their formation (“primary
difl'erentiation") and the subsequent processes of histologiml and
morphological difierentiation. At the same time the mammalian egg
was at last discovered, and before long recognised as a single cell.‘ The
nucleus of the egg-cell had been seen in molluscs as long ago as t79x
by Poli in Xtaly, but not cleaxly dscribed until the xwtk of the Czech
Purkyne on the "germinal vesicle” of the hen's egg in x825. In this my
the road lay open for the triumphs of the mid-nineteenth century, when
a Kowalevsky could reveal (1867) that such fundamental processes as
germ-layer formation and gnstrulation were common to all animal
phyla.‘
 
It is interesting to note, however, that the recapitulation theory, which
was first clearly formulated by Von Baer, was already taking shape in ~’
various minds during the closing years of the eighteenth century.‘
Lewes has thus described the thesis of Goethe’s Morplxologie, written in
 
W957
 
The more imperfect a being is the more do its individual parts resemble
each other and the more do these parts resemble the whole. The more perfect
a being is the more dissitnilarare its pam. In the former case the pun are
more or less a repetition of the whole, in the latter case they are totally unlflte
the whole. The more the parts resemble each other the lm subordination is
there of one to the other: and subordination is the mark of high grade of
 
organisation.‘
“’i|liarn" and John Hunter belong also to the end of the oentury.
'-‘For ‘xniumuftion an ‘van But. he Ks:-stv, Atddtfln; A. “I. Maya’. Ind. Stink-
 
h ‘ ( ' t z. _
H)I,fI“:$>.i‘s°gu:i>e hii:°mg?§Esms:§n.m's work on the development of Amphibxl (J.
Pagel).
 
11,,-, k {S h ' 8 Cf. E. 5. Russell. A
 
: c[_D‘g‘;!d:;rieO‘p'snplenl‘;ein1cet'w;|-glrélr|I?\x1ss3e<ll;md the classical mm at;-‘mt muoux.
 
‘ The history of the coneepraf rccspmzhuan has recently been gone mm with name
thoroughness by A. w. Meyer (I93s).who is ttr-nzely rel!-Icunt '0 511*‘ '“Y6“'““
it earlier than Hunter. On the other lnnd, I am glad to Ice thxt Bxlsl (Iv: ) IS?‘
with me ugnausg Ammo: on this matter (m p. 49 of the vmcm hm‘)-
 
- L,-/-,, F, 353, ' 5:: Duncan.
 
224
CHART III
 
 
1450
 
CHARTTO
 
ILUJSTRJNTE. THE. Hl5TOR,Y
(if EM
 
ranvomcv mom 1920
a.d.to1p10u.d..
 
1600
 
2:5
 
n.2.—:5
A HISTORY OF EMERYOLOGY
 
‘_2‘;ll°fi?:1€F. In his book on the anatomy of the gravid uterus, proved
_ y an completely the (l'uEh‘0f the new that the maternal and foetal
circulations are distinct. HIS injections left no shadow of doubt about
ll"? lTlIll_1€l'. and the way was clearly opened up {or the study of am
Pl’°P€l'tI¢8 of the Capillary endotheliztl membranes separating the bloods.
a study which is still vigorously proceeding both in its histological and
physroco-ehemicnl aspects. There was a quarrel between thehrothers over
the priority of this dernonstmtion. John Hunter's Essays and Observa-
tion: also contain matenal important for embryology.‘ His drawings of
the chick in the egg were very beautiful, and are still in the archives of
the Royal College of Surgeons. He adopted l\iayow‘s theory of the
oflice of the air-space, and anticipated von Baer’s theory of recapitula-
tion much as did Goethe.
 
If we were capable of following the progress of inurease of the number of
parts of the most perfect animal as they were first formed in auccesion, from
the very first to its state of full perfection, we should probably be able to
compare it with some one of the intxztnplete animals themselves. of every
order of animals in the creation, being at no stage difierent from some of
the inferior orders. Or in other words, ifme were l0 talic 3 "55 05 an-lm3l3.
from the more imperfect to the perfect, we should probably find an imperfect
animal oorruponding with some stage of the most perfect.
 
It is interesting to reflect on the curious course which was taken by
the essence of the idea of recapitulation in the history of embryology.
As first formulated by Aristotle it was as much bodily 3-5 mental» but all
his successors until the eighteenth century treated it as a psycho-
logical rather than a physiological or m0rphol0glC3l “~ll=°TY. and lit“
themselves in speculations about the vegetativc. Senslllvc and rational
souls. Yet the other aspect of the theory was only aslctp. and W35 des-
tined to be of great value as soon as investigators began to direct their
attention more to the material than to the psyclmlogitzl aspect of tilt
developing organism.
 
Hunter did not absolutely reject preformationlsmi ll‘-ll l’¢Ef1“l¢l'l ll 3-‘
holding good for some species in the animal lilflgdflmi l1= ll1=1’¢f°Y°
attached no philosophical importance to it.
 
Although “'olfl"s work did not lead to the immediate morphological
advances which might have been cgpected. it was in mm’ "t"-‘Y5 {"‘“‘f“l-
It stimulated J. F. Blumenbaclfs Ubrr Jen Bildifllglhitb Of 1739. fl_“’0fl§
which elaborated the Wolfiian tr‘: trrenlialir into tl1t=_III'-Y1l3[0f7fWl!’|lf. 1*
directing morphogenetic force peculiar to living bodies. ll 15 lnlefcfilmg
I Detailed and interesting Iurveys of John HuntefIdc°I:l:v1;l:UllIll":"-"dl;ed"‘;'g"3"E£°g,7
 
have been published by A. W. Meyer (1915. 1916). In
his address at the Hnrnrd Bicentenary Celebi-mom of EM!“-
 
226
EMBRYOLOGY IN THE EIGHTEENTH CENTURY
 
to note that Blumenbaeh passed through an'exactly opposite succession
of opinions to that of Haller, for he was first attracted by preforma-
tionisrn, but being convinced by VVolfl"s work,‘ abandoned it in favour
of epigenesis. Blumenbach comparu his ninufarmativur with the force
of gravity, regarding them as exactly similar conceptions and using
them simply as definitions of a force whose constant efiecrs are re-
cognised in everyday experience. Blumenbach says that his nimr
farmativur differs from \Volfi’s vi: errmlialxlr because it actively does the
shaping and does not merely add suitable material from time to time to
a heap of material which is already engaged in shaping itself. Woltfwas
still alive at this time, but made no comment on Blumenbach’s ideas.
He may well have thought the differences unimportant. Both Blumen-
bach and Wolf? were mentioned by Kant in the Critique of jiuigment,
where he adopted the epigenetic theory in his discussion of embryonic
development.
A word must be said at this point about the opinions of the eighteenth
century on foetal nutrition. At the beginning of it there was, as has
been shown, a welter of conflicting theories, and though later on writers
on this subject were fewer, the progress made was no more rapid.
In 1802 Lobstein was supporting the view (which had been defended
by Boerhaave‘) that the amniotic liquid nourished the embryo per us,
though Themel had shown forty years before from a study of acephalic
monsters that this could be at most the very slightest source of material.
These workers had obviously learnt nothing from Herissant and
Brady, who had been over precisely the same ground fifty years earlier.
On the other hand, Good and Osiander reported the birth of embryos
without umbilical cords, so that the solution of this question became,
in the first years of the nineteenth century, balanced, as it were, be~
tween the relative credibility of two kinds of prodigy. Nourishment
per 0: was defended by Kessel, Hermes and Grarnbs, and was attacked
by Vogel, Bernhard, Glaser, Hannhard and Reinhard. The idea lingered
on right into the modern period, and as late as 1886 von Otr, who was
much puzzled about placental permeability, decided that a great part in
foetal nutrition must he played by the amniotic liquid. Weidlich, a
student of his, fed a calf on amniotic liquid for some days, and as it
seemed to thrive on this diet reported the amniotic liquid to have
nutritive properties. The appeal to monsters was still resorted to at the
end of the nineteenth century, for Opitz, in order to negative Von Ott's
conclusions, drew attention to a specimen in the Chemnitz Polyklinik
in which the oesophagus of a well-nourished normal infant was closed
 
' And by experiments of his own on the regeneration of hydroidr.
‘ fInm‘nm'on¢:. net. 382.
 
2 27
A msronv or cxuanrocoor
 
at the upper third without the development of the body having been in
any way restricted. The fuller possibilities of biochemistry itself have
sometimes been exploited in favour of the ancient theory of nourish.
mm‘ Per 0!: thus Kfittnitz in 1889 collected some data about the pres-
ence of peptones and protein in the human amniotic liquid with this
object in view. That the foetus swallows the liquid which surrounds it
towards the end of gestation in all anmiota can hardly be disputed, and
as there are known to be active proteolytic enzymes in the intestinal
tract, no doubt some of the protein which it contains is digcsted—but
to maintain that any significant part is played in foetal nutrition by this
process, has become steadily more and more impossible since 1600.
 
But to return to the eighteenth century; all was not repetition,
occasionally someone brought forward a few facts. Thus the degluti-
tion of the amniotic liquid was discussed by Flemying in 1755 in a
paper under the title "Some observations proving that the foetus is in
part nourished by the amniotic liquor." “I believe,” he said, "that very
few, if any at all, will maintain now-a-days with Claudius de la Courvée
and Stalpart van-der-Wiel, that the whole of its nourishment is Con-
veyed by the mouth." But he himself had found white hairs in the
meconiurn of a mlf embryo with a white hide. Both Aides and Swam-
merdam had found the same thing. but Aldes did not think it of any
significance, and Swamrnerdam merely remarked that the calf must
lick itself in utero.
 
More interesting was W. Watson’s "Some accounts of the foetus in
ultra being diflerently affected by the Small Pox.” This was the earliest
investigation of the permeability of the placenta to pathological agents.
“'I'hat the foetus," said Watson, "does not always partake of the _Infec-
tion from its Mother, or the Mother from the Foetus, is the subject of
 
this paper.” Two of his cases, he said,
 
evince that the Child before its Birth, though closely defended frorn the
external Air, and enveloped by Fluids and Membranes of its own, 15 not
secure from the variolous Infection, though its Mot.her.has had the D15-
temper before. They demonstrate also the very great Subtilrty of the vanolous
 
Etfluvia.
But other cases
 
are the very reverse of the former, where though from inoculationthe mgst
minute portion of Lint mo1sten’d with thc\:anolous Matter and _5PP]_‘°d ‘°
slightly wounded Skin, is generally sufficient to pmpngyite this D1Sl_¢'“P¢1’u
yet here we see the whole Mass of the Mother’; Blood. urcglaung dunno the
Distemper through the Child, was not sulhuent to produce it. . . .
Historic: it appears that the Child before IE mnh ought to 5* mm“ “
 
228
EMBRYOLOGY IN THE EIGHTEENTH CENTURY
 
separate, distinct Organization; and that though wholly nourish'd by the
Mother’s Fluids, with regard to the Small Pox, it is liable to be affected i.n a
very different Manner and at a very different Time from its Mother.
 
Doubtless the modern explanation of \Vatson’s discordant results
would be that in one case there were placental lesions, destroying the
perfect barrier between the circulations, and in others there were not.
In the last year of the century (but the seventh of the Republic)
Citizens Lévcillé 8: Pnrmentier contributed an interesting paper to the
journal de Physique in which they observed the increase in size of the
avian yolk an incubation and spoke of a current of water yolkwards.
CONCLUSION
 
WI_IBN the contents of this book was given in the form of lecturu at
University College, it bore the title “Speculation, Obsenvagion and
Experiment as illustrated by the History of Embryology.” Of the fin:
two of these factors we have seen enough, but the third would have
necessitated the continuation of the story down to the end of the nine-
teenth century, and this must still await our projected second ‘-0113,15
The true science of experimental embryology did not come into being
until the time of Wilhelm Roux} The early chemical Obsefijfions an
the embryonic liquors (p. r 59) were indeed observations rather than
experiments, and there was no systematic study of the changes which
the liquors undergo during the development of the foetus; this was
not done till the time of John Dzondi (1806). Harvey’s segregation of
does at Hampton Court (p. r46) merits perhaps the name of an experi-
ment, involving as it did die use of "cantrols;" and an outstanding
instance is the ligature of Nuck in x691 (p. 163). As in Nuck‘s case,
experiment in the hands of bath Spallanzani and J. T. Needham led to
error. Spallanzani confuted his adversary on the question of spontaneous
generation and the vegetative force by what amounted to rigid criticism
of experimental conditions, but later on denied their proper function to
the spermatozoa on exactly the same methodologically faulty grounds.
But on the whole, experimentation, active interference with the wurse
of Nature and subsequent observation of the resulting system in
comparison with systems in which no such interference has taken place.
was a characteristically nineteenth-century product as far as biology and
embryology are concerned. Only at the prrsent day, indeed, are rte
beginning to appreciate the statistical and other difliculties attending
upon the full applimtion of the experimental method to living organ-
isms, and the manifold obstacles which prevent obedience to the rule
that only one variable be modified at one time. But this is no matter of
reproach against the older embryologists. Knowledge of form must
neoessarilypreoede knowledge of change of fon-n and the factors pro~
ducing it, and so we see during the last seventy years the production of
 
Info]. Entu-'ckbmgrm:z}:4nilr in 1394. thus naming
ta! morphology and arm! embryology.
 
23o
 
‘ 185°-tgu. He founded the Art
the modern discipline of erpenmen
CONCLUSION
 
"Nomialtafeln" or tables of morphological pictures showing normal
development; these are the essential basis for experimental studies.
 
Probably the best way to summarise the influences which have
operated in the history of embryology is to concentrate attention on
what may be called, borrowing a phrase from general physiologifi-the‘
"Limiting Factors” of advance. We may thus regard the progress of
knowledge about generation as governed by a reaction-chain, one link
in which may at any given time be slower than all the others, and hence
may set the speed for the whole.
 
Relation of lnvesrigarnn to their environmmt
 
Co-operation of investigators
 
Prminl
mm“: {mm recrmique<n . Terminologzcll C
Wftufli UAIRUSGVE
Cancepruzl {
Destructive
 
Psychologies!
 
Balance berween Speculation, Observation and Experiment
 
Of these limiting factors the first which may be mentioned (though I
do not wish to pronounce here upon their relative importance) is the
relation of investigators to their environment. The Carlylean tendency
 
to regard the history of science as a succession of inexplicable geniuses ‘
 
arbitrarily bestowing knowledge upon mankind has now been generally
given up as quite mythological. A scientific worker is neoasarily the
child of his time and the inheritor of the thought of many generations.
But the study of his environment and its conditioning power may be
carried on from more than one point of view. We have already seen
(p. H 5) what a sharp distinction the culture-historians (Sigerist, Bili-
kiewicz, etc.) make between the mental atmosphere of the Renaissance,
the Baroque, the Rococo, the "Au£‘k!.amng" period, and so on. There
is doubtless much to be learnt from historical investigations carried on
in this light, but it may sometimes lead to a hypostatisation of abstrac-
tions, and as in the case of ovism and feminism (p. H6) its results may
border on the fantastic. The social and political ruling ideas of a dis-
tinguishable epoch play, on this view, an overwhelmingly important
part in the scientific thought of the time, and may act as limiting factors
to further advance. Thus the politiml absolutism of the Baroque period
is thought to have mirrored itself in the extreme rationalism of seven-
teenth-century biology. There is much more to be done in working out
 
23:
A HISTORY OF EMDRYOLDGY
 
the internal relations of successive intellectual climates and their con-
nections with contemporary social situations.
 
The other principal point of view which may be taken regarding the
environment of the scientific worker as a limiting factor is that which
emphasises his existence as an eccnoniicunit, andscelts to show horrhis
position in a society with such and such a class structure influences the
development of his thought. Some reference to this point of View has
already been made in the introduction (p. r4). It seems to ofl'er more
chance than the preceding method for new discoveries in the history
of science, for it directs its attention upon those aspects of human
society (trades and techniques, labour conditions, the evcryda ' life of
the mine, the factory, the barber-surgeorfs shop) which, precisely
because of their assumed inferiority, have not been discussed in the
majority of books, written inevitably by members of the goveming
classes, or by those who aspired to imitate gentility. Thus the rather
sharp cleavage between the philosophic biologist of the Hellenistic age
and the contemporary medical man, who might often be a slave._c_on-
tributed doubtlss to the sterility of ancient Mediterranean x;nCdlClIlC,
including obstetrii: and gynaecology. ln the later Christian \\ est there
was not much incentive to embryological study so long as the process of
childbirth was left to the charms and incantations oi: barbarous nud-
wives. But for a better insigbl l“‘° am °°°’,‘°m‘° P‘-"5‘"°“ °f °mb’5’°'
iogiscs in past ages nearly all the work ri-mains to bedpne. .
 
Next Domes co-operation of scholars. In the civilisation of the
Hellenistic age, it may ‘be said, a considerable measure af_ such 43*
operation had been attained;.the iivorks of Aristotle and Hipgsocss as
were fairly readily available in iyritten form, and evidence CE:
brought {onward (pp. 54: 73>. pm-cuiulywuh r=sardt°J=“r1sb "*g.“§ ~
that this was well used. But we must beware here of sufienng. a is ctr‘;
(ion of perspective in the contemplation of antiquity,_lu1l’ It’: ¢35Yu1d
exaggerate the co-operation of ancient thought A 5“‘l:' '5 ‘ :‘c::cm
consider itself lucky if it passed once in twenty-EVE 1:313 fljcfin
Greece and India after Alexander  13- 27)- Am°"E  °°‘t'ms B:
influences that save fis= *° ‘‘*° '-‘“i“'”?'°.“ "5 "“ ‘-M flinehanci
aperation,hamperedbyenoimouslmgltlsfltf d1fli°‘1l“_53c;“ dihcolo _ul
and by the diversion of interest £rom scientific to etlhrl 311 C mVc3‘!hc
channels on the other, sank to a V60’ 1°" l°‘°l' ah°‘:l°°of‘;is wmem_
|’¢m5“'k“b1° Specude uf 2 Leqnirdo’ many Y?’ Z of fortifiutlonsi
pomrics’ and able to am 3 lmngfmly all adimgriries to any living
finding it impossible to ooxnmmucate  Ksgovks my by 3 mm
person. and reduced to 5“-YY“‘§ am“ “‘ “° C M
chance available to SC.l‘tOl£il'S of after 3855-
 
232
CONCLUSION
 
Among the most important of limiting factors we must reckon
technique, extending the term to cover mental as well as material
methodology. The part which the latter has played in the history of
embryology can hardly be overrated. Thus until the introduction of
hardening agents, especially alcohol, by Boyle  158), the examination
of the early stages of embryos was bound to remain crude, and we have
seen (p. 185) how embryology attained an entirely different level im-
mediately afterwards in the hands of Maitre-Jan. The parallel case of
the microscope is too familiar to dwell on, but the work of Malpighi
obviously marked a turning-point in the science (p. r63). It may here
be noted, however, that even when methods are available, the workers
of the time do not necessarily use them, and although Harvey could have
employed an early {am of microscope, he restricted himself to the
weak lenses, pnrpfdlia, or "perspectives,” which had already been
used by Riolanus.‘ A still more striking instance is that of artificial
incubation. Carried on in Egypt since the remotest antiquity (p. 22),
this process must have been at the disposal of Egyptian physicians,
Alexandrian biologists and Arabic scholars for a period of three
thousand years, yet so far as we know, no embryological use of it was
ever made. In eighteenth-century France and England the technique of
the process had to be painfully rediscovered at a time when biologists
were only too eager to make use of such assistance. Let us mention, as
other instances of the effect of material technique on embryology, the
burst of knowledge which followed the invention of the automatic
microtorne by Thrclfall and others about 1883, and the great advance
which in our own century has followed the successful mastery of graft-
ing technique in operations on amphibian embryos by Gustav Born and
Hans Spemann.
 
Just as important, however, as material technique is mental technique.
And first with respect to words; on several occasions we have had to
notice a standstill on account of the lack of a satisfactory terminology.
 
‘ As this rtzrterxrent lbout Harvey may seem Iurprising to lame, it is worth while to
rcopitulate briefly the fact: thou! the iruention of the microscope. For the detailed
tvidenee the elassrml papers of Singer may be consulted. The introduction of convex
lenses Is spectacle: for presbyopin may now be dated very soon after A1). 1286 in
Iuly, recording to the ¢X.hl|.I3LlV€ researches ol Roscn. Concave lenses for rnyopin
came into use much later, about xsoo, Ind the first bilcnticuhr ryutem was probably
due to Leonard Digger (d. r57r), who invented (but never fully described) I rudimen-
tzry telescope. The Eu-rt lens combination for a microscope Wu mentioned, again
obxcurcly, by Gin:-nblttistx dell: Port: (1540-1615) in his Jllagiae Nannalix (1559),
but no practial nppliution was made cm. The rnicroscope mu, begituwith bclurias
Jansen (I530-fa. 1630) of Middclburg in Holland, who put together two convex
lenses rome time between 1592 and recs, as we know from the detailed Ineount of
Pierre Borcl. Comoliu: Drebbel of Alcrnur, nuthcxmticiln to King June! in X619.
brought one or these instruments into England. i e. thirty year: before the conclusion
 
of Wrtlrun Haney’: researches on genennon. Perhaps Harvey made :Eom to acquire
and me one, perhaps he was too uonren-nL~i-V: and letpticaI—'we do not know.
 
333
A nrs-roar or zrmarotocr
 
Thus in the thirteenth century Albertus of Cologne had arrived at a
Pom: bcwnd “inch P1987335 was impossible in the absgnee 1‘
words’ when’ {°' ‘“-mPl°o ‘hut Was no other means of descrihingnte}:
sero-amniotic ’unction ’ t c h ' - ..
 
on the left sidje of the i’-essigl whicherizghsllhébloiig  bihn«:hi: htfiic
right hand of something else” accuracy was difficult and s eed in‘: C
aim“ A P795551)’ Sirnilar position was occupied by Boerlliaave
Clflhtcfimh 591""-1'Y. only now in the case of biochemical word; Factd
“-kh.s°m° 5"b5“"“°° sud‘ 35 3 "EYCOSY, streaky yellow oil, smelling of
3u“111n¢ 531$." Boerhaave  unable to describe it except in thsc corn.
m°"‘5_¢D5° "M15: and lacking the means either to submit it to further
analysts or to characterise it by accurate physiCo<.hcmiml units, he was
forced to admit a large number of ultimates into his schemes which
were not ultimate at all.
 
Mental technique as a limiting factor in embryologiml history goes
deeper than words, however, for it involves the concepts of the investi-
gator. What the Germans mil “Begrifl"sbildung" or the construction of
concepts congruent with oertain sorts of natural phenomena. though
never conscious in the history of biology, has none the l$s been open.
tive. In this field we may remember the doctrine of Galen conoeming
the natural faculties (61-rci;:s1;. p. 70), and the immense length of time
which was required for biologisw to see that it was nothing more than
8 concise statement of the phenomena themselvms. Not until it was
"seen throng " as an explanation was post-Renaissance biology pas.
sible. Similarly, the peculiar contribution of Leonardo to embryology
was his realisation that embryos could be measured. not merely as to
dimensions at one moment but as to dimensions at a suocasion of
moments. The applicxtion of the concept of change in weight and size
with time, a concept which, as modern biology $hO\\S, admits of much
accuracy when properly worked out, was thus first made by Leonardo.
In the same way Boyle was the first to see clearly that a problem of
mixture is presented by the developing embryo (though Hippoa-ans had
stated it dimly some two thousand years before). If the embryo is made
up of mixed things, some definite proportion and way of mixture rrnrst
exist. And no hope of finding out what this was could be obtained from
the Aristotelian elements (heat, cold, moisture and dryness) or from the
alchemical principles (salt, sulphur and mercury). Hence Boyle’:
emphasis on the corpuscularian or mechanical hypothais. and all its
 
historical implications (p. 176).
Besides this creation of concepts, and the choice of which of them
to apply, the mentality of the scientific workers of the past often
 
difiered greatly with regard to a fundamental quality which can WIY 5'?
234
CONCLUSION
 
called audacity. Probably Aristotle's greatest claim to our respect is
that alone of his contemporaries and predecessors he had the audacity
to suggest that animal farm is not limitlessly manifold or infinite in its
manifestations, but that given industry and intelligence, a clzssifizmtion
was possible. This alone marks him out above all subsequent biologists.
On a smaller scale, we find the same mental audacity in Kenelm Digby,
whose discussions of the development of the chick are remarkable for
their naturalistic tone (p. 122), for their conviction that the proccssu
of development are not beyond the reach of the reason and imagination
of man. It is most ironic that Digby, who did little or nothing himself
to advance our knowledge, should have spoken thus, while his great
contemporary, \Villiam Harvey, to whom we are indebted for so many
advances in embryology, was led to despair of understanding develop-
ment. Another interesting point that emerges from the same period is
that such mental audacity can go, perhaps, too far, as when Descartes
and Gusendi built up an embryology more geometiico demomtram, in
which the facts were relegated to an inferior position and the theory
was all.
 
But not only must the right concepts be chosen, the wrong ones must
be abandoned. One of the principal necessities which has faced investi-
gators since the earliest times has been the recognition of silly questions
in order to leave time for the examination of serious ones. It was
presumably inevitable that the pseudo-problems concerning the entry
of the soul into the embryo should be taken seriously until a very late
date. But a more typical instance of a meaningless question may be
found in the dispute about what parts of the egg farm the chick and
which feed it. The tacit assumption here was that since to common-
sense food and flesh are different things, there must be in the hen’s egg.
bsides sufficient provision of food, some sort of pre-flesh out of which
the embryo can be made. Not until 1651 did this pseudo-problem go
out of currency in the light of Harvey's demonstration of the unsound-
ncss of the assumption.
 
The expulsion of ethics from biology and embryology forms another
excellent example. That good and bad, noble and ignoble, beautiful and
ugly, honourable and dishonourablc. are not tenns with biological
meaning. is a proposition which it has taken many centuries for bio-
logists to realise.
 
Idus of good and bad entered biology partly under the concept of
“perfection? In 1260 Albcmrs was maintaining that male chicks
alvay: hatched from the more spherical eggs and female chicks from the
more oval ones, because the sphere is the most perfect of all figures in ’
solid geometry. and the male the more perfect of the two sexes (p. 87).
 
=35
A HISTORY or XZMBRYOLOGY
 
‘WC realise today that to ask which is the more perfect of the two sexes
18 1| meaningless question, for we have expelled ethics from science and
°=’“}°‘ |’=Lr~1fd any one thing as being more perfect than anything else.
A831": describing the course of the arteries in the developing chick
 
Mb"-""5 “F53 "One of the two pusagm uhich spring from the heart,
b"1"°hC3 info ‘W0. One of them going to the spiritual part which contains
'-he ha“. and carrying to it the pulse and subtle blood from which the
lungs afldotherspiritual parts are formed; and the other passing through
the diaphragm to enclose the yolk of the egg, around which it forms the
liter and stomach." This distinction between the organs above the
dflphmgfn. the lungs, heart. thymus, ete., cxlled "spit-itualia,” and
the organs below, the stomach, liver, intestines, spleen, ete., runs
through the whole of the early anatomy. It was as if the organs of the
thorax were regarded as a respectable family living at the top of an
otherwise disreputable block of flats. To us it seems absurd to call one
organ more “spiritual" than another, but that is because we realise
the irreletance of ethic-:.l issues in biology. Thomas Aquinas, about
the same time, dealt in passing in his Summa 77zealagz'm with human
generation (p. 93).
 
The generative power of the female is imperfect compared to that of the
male; forjust as in the crafts, the inferior workman prepares the material and
the more skilled operator shapes it, so likewise the female generative virtue
provides the substance but the active male virtue makes it into the finished
product.
 
This is really the pure Aristotelian doctrine, but St Thomas gives it the
characteristically mediaeval twist. Aristotle might make a distinction
between form and matter in generation, but the mediaeval mind, with
its perpetual hankering after value, would at once enquire which of the
two, male or female, was the higher, the nohler, the more honourable.
 
In the eighteenth century the same frame of mind persisted. It was
maintained that in every detail of the visible world some evidence could
be found for the central dogma of natural religion, the belief in 3 .l“33
and beneficertt God. Biology was thus not free from the mental [>515
associated with theology.‘ Between 17:20 and 1850 a multitude of books
were written which purported to reveal the wisdom and goodness of
 
God in the natural creation. The theologians tool: what suited their
purpose and left the rest. It is instructive to see how Goethe. who was
deeply committed to the theologiml interpretation of phenomena, re-
acted to the ornithological anecdotes of his secretary E'cke1-mano on
18th 0et., x827. He said little while Eckerrnann told him about the
 
1 Far I ‘pg-[king mmple of this. see Edmund Gone‘: I-‘atlnn and San.
236
CONCLUSION
 
habits of the cuckoo and other birds, but when Eckerrnann related how
he had liberated a young wren near a robin‘s nest and how he had found
it subsequently being fed by the robins, Goethe exclaimed:
 
That is one of the best ornithological stories I have ever heard. I drink
success to you and your investigations. Whoever hears that, and does not
believe in God, will not be aided by Moses and the prophets. That is what I
call the ornnipresence of the Deity, who has everywhere spread and implanted
:1 portion of His endless love.
 
And so it always was with the theological naturalists; they hailed with
enthusiasm the discovery of monogamy in tortoises, or mother-love in
goats,‘ but they had nothing to say concerning the habits of the hook-
worrn parasite‘ or the appearance of embryonic monsters in man. Not
until the beginning of the nineteenth century did it become clear that
Nature cannot be divided into the Edifying, which may with pleasure
be published, and the Unedifying, which must be kept in obscurity.
Experimental embryology then contributed to this clearer vision of the
living world by its manifold demonstrations that in spite of the appar-
ently deeply teleologiml character of normal embryonic development,
once the individual morphogenetic processes have been experimentally
"derailed," they laboriously continue their operations so as to imitate
(and therefore ultimately to explain) all the possible varieties of naturally
occurring monstrosities.' Of course the riddle of nomml integration
remains.
 
In the end we may say that the progress of a branch of natural science
such as embryology depends on a delicate balance of three things,
speculative thought, accurate observation and controlled experiment.
Any modification of the optimum balance will act as a powerful limiting
factor on progress. Speculative thought, in particular, has shown a
tendency to crystallise too readily into doctrines which, by way of
attachment to some philosophicafor theological issue, live a longer life
than they deserve. Thus the Aristotelian theory of the {emotion of the
embryo by the coagulation of the menstrual blood, built in the first
 
‘ One find] stfilting parallels for this interest in animal behaviour among the neo-
Cornfucian school of philosopher! in mediaevnl China (see Sciznzz and Civtlirntion in
China, vol. a). In so far as it contributed to n usnviction of the reality of an evolutionary
 
ITKJCSS. which for the Chinese thinkers it certainly did, it was useful and commendable.
 
ut that-uheywere never committed to the idea of a rpeciat creation by uuu-benzfieent
personal deity. For them thertforc "gleam: of righteousness” in ants and alien were
pres: of that human community which the im enonal Order of Nature (the Tao)
wool in due time produce, pieces of evidence 1 out 3 social evolution, not about a
personal Creator.
 
‘The guinea-worrrI(Dra.I.1mnlIu.r irmimemis) had been givzn . drunatic d:_scrjption
by Vzlsch in r67.;—nnd indeed by Avicenna long before him. A.n.Lylostorru1ns had
been known Ind described in ancient Egypt.
 
‘ On lethal genes and their action, see the brilliant book of Hxdorn.
 
337
A HISTORY OF FMBRYOLOGY
 
instance upon a faulty deduction, became incorporated in the Aristote-
lian tradition of form: and materia, and although quite repugnant to ob-
servation, remained the official theory throughout the European Middle
Ages, and apparently  perpetuity in India. So ponerful was the
rationalism of a medical education round about 1639 that the physicians
to whom Harvey demonstrated the empty uteri of the king's does
preferred to believe their books rather than the evidence of their senses.
And precisely parallel to this attitude, as we have seen (p. 213),  that
of the preforrnationists in the following century, who, having dccfl‘-ltd.
like Bonnet, that epigenesis was inconceivable, only aooepted such
observations as coiilinned their upriari view.
 
Prefomiationisin as a manifestation of rationality merits’ further
examination. The dogmatic manner in which preforinaticnism W-is
held during the eighteenth century would not perhaps have bceit 50
fatal if the biologists of that time had been able to take rnatheniatical
reasoning more seriously. There um Harvey's yery convincing ai‘B“'
inent about the circulation of the blood, and Freind's equally convinc-
ing. but unfortunately erroneous, deductions about the quantity of
menstrual blood and the weight of the newborn foetus (p. 1'50). If these
could have been accepted, it was a pity that I-Iartsoekers argurnglit
about preforrriation could not. In 1722 Hartsoeker caleulated t I
l0‘°°'°°° rabbits must have existed in the first rabbit, 35-Shummg '11“ ‘he
creation took place 6000 5'63” 33° and lb“ }"’bb"s bggm la "Pmduc;
their kind at the age of l‘KblOnlll1'|Sd;'nBUt to I135 B‘J}:sn€tY°"c1:“"3)¥'
thatitwasalwayspossi 6, Y3 33"“ ‘’““f ' . '
nation under the weight of numbers, and he described !h= P"5‘{"m3“°“
theory as one of the most striking victories oi the understa:_u‘g:Ig 0::
the senses. It would have been better dumbed as one 0 9 In
striking victories of the imagination over the understanding. . d ‘E
 
The fact is that the biologists of the eighteenth century, cxirne Jar:
by Prefoririationist theory, tool: emb1'Y°l°EY °“ ‘"3 Pmteagllc ‘he
observation became superfluous. They rrould have founm alieepd hm’.
sentiment satiriscd by Boyle that “ us.  mote E 3:11" P do‘
sophiczil to argue a prion" than 1'1?”-"”“’”' and min ewilidionythnt
barred from looking at devclopmg €mbl_'}‘03 by their con‘: ‘her may
structure and organisation would Ifertflilili 59 ‘l‘°‘°' W cm {ad I
'°°"’d 5°‘ i‘ °‘ “°“ Th‘ "'°{°'"“"°"’“ mwowtify liiiaibhalists {mi
“P5550” in bi°l°gY °f ‘he mnufltfly between C rtionalists were
the ernpiricists in pliil0S°PhY- The wnmmpomry ta
 
Pwpk who held that - - - i f’ mpmatien which
humwI>=ina= "=r°i*=P‘zss°?“°“°‘°‘"““ ‘3“"”*’ “° “‘ d aids. be
were not simply g€fl€1'IlJSal.IOEB from expencntfi, 5“ W“, "C"
 
238
CONCLUSION
 
used as major premises in arguments concerning Nature. If observations weie
not in accordance with expectations founded on such reasoning, they were
dismissed as illusions. The empiricists, on the other hand, held that there
was no knowledge independent of observation, and that the rationalism’
principles, in so far as they were admissible at all, were generalizations from
experience.‘
 
It is obvious that nearly all the preformationists were rationalists. They
thought that Reason was in a position to decide the issue whatever
might be the results of observation. "It is remarkable,” as Cole says,
in his book on this period, “that the preformationists did not realise
that if the point to be established is assumed at the outset all further
discussion is superfluous.” In this example, then, we have a disturbance
of the balance towards the side of rationalistic speculation.
 
It would be a mistake, however, to regard this tendency as confined
to the eighteenth century. Ample examples of its presence can be
collected from nearly every period in biological history. “We plume
ourselves,” says Cole, “on that aspect of our work which is vain and
argumentative, and condescend to the more modest but enduring
labour of observation." There can be no doubt that this state of afiair-s,
so unfortunate for science. is one aspect of that contempt for manual
labour which has run through the Stratified structures of all societies in
the history of civilisation. The manipulator of paper and ink, educated
in the classical traditions of his time, has always seemed, by reason of
his superficial similarity to the political administrator, a superior being
to the empirical mechanic engaged in the manual work of the arts and
industries. The tradition is as old as civilisation, yet for the advance of
science it must be broken. Not until the manual worker and the
audacious theorist are combined in one person will the fullest develop-
ment of scientific thought be possible.
 
On the other hand, there can be no doubt that a plethora of observa-
tion and experiment is also had for scientific progress. Modern biology
is the crowning instance of this fact. What has been well called a
 
"medley of ad hoc hypotheses" is all that we have to show as the
theoretical background of a vast and constantly increasing mass of
observations and experiments. Embryology in particular has been
theoretically dxraidbare since the decay of the evolution theory as a
mode of explanation. Embryologists of the school of F. M. Balfour
thought that their task was accomplished when they had traced a
maximum number of evolutionary analogies in the development of an
animal. \ViIhelm His, perhaps the first causal embryologist, struggled
succtssfully to end this state of afi'airs.
 
I \\'oodgtr.
239
A HISTORY OF EMBRYOLOGY
 
My own attetrrpts [he wrote in r888 in 2 famous passage] to introduce mm:
elementary physiological or mechanical explanatinns inta ernbryolog have
not been generally agreed to by mnrphnlogisu. To one it seamed ridiculous
to speak of the elasticity of the germinal layers; anatherrbought that by such
considerations we put the can befure the horse; and one recent author states
that we have something better to do in embryology than to discuss temions
of germinal layers, etc., since all ernbryologiml explanation must nccaxtrily
be of a phylogenetir: nature.
 
But this strictly evolutionary dominance in embryology did not last on
into the twentieth century. The unfortunate thing is that nothing has
so far been devised to put in its place. Etperimental embryology,
Morplrologiml embryology, Physialogical cmbryology, and Chemical
embryology {am today a vast range of {actual knowledge, without one
single unifying hypothesis, for we cannot yet dignify the axial gradient
doctrines, the field theorim and the speculations an the genetic control
of enzymes, with such a position. We cannot doubt that the most
urgent need of modern embryology is a sex-res of advances of a purely
thearetitzl, even matlrerrrntiw-logical, nature. Only by samething of
this kind can we redre§ the balance which has fallen over to observation
and experiment; only by some such cflort an We obtain a theoretrral
embryology suited in magnitude and spaciousrtess to the wealth of facts
which contemporary investigatnrs are accumulating day by dd)’-
 
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253
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276
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Waddingmn. C. H. T11: Epigmettle: o/Birdx. (C.-unbridgt, 1952.]
Waddingtan, C. H. Pn':ta:1:I:.r aj'EmbryaIogy. (Allen .1: Umvin, London. xg56.)
Waldschmidt. J’. Pmxir Illedirinae Rationalix, pp. 722, 743. 744. (Paris, 1691.)
V'v':|XI:r, R. Phil. Tram. Roy. Soc. 1693, x7. 523.
 
van Wasscrberg. L. Bald'x'ng:r': llfagazfn. 1780, 2, 300.
 
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Wcinxich, Martin. D: Ortu lllorulronnn Commentariru, in qua mtntia, '
 
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(Breshu (T). 1595-) _ _
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39°
BXBLIOGRAPHY
 
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\VurcesLer, Henry Samerset, 1st Marquis of. A Century :7] the Name: and
Srzmllings of such Invention: as at present I can tall to mind to hem tried.
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291
A HISTORY OF EMBRYOLOGY
 
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’ 1929. xx. 55.
 
Zypacus, F. Fundanunta mdka. (Bmssds, x684.) -
 
The following work remain in-accasihla to me in Englfind 1nd 1 MW 110‘
yet seen them:
Mzntcllassi, c: Dr'vm1'S:'.rmm':an L: Gtnzraziane. (norcncc, X749-) Rvpfimd
in Rama’! dz pike: tie Illldez-in: (Paris, 1763) and in Pféa: inllrmdnfef
:10 la Mldea'rze. (Luny. Pan's, 1782.)
1: Monnier, P. D: Conzepm ct Irmzmenta Foetu. (Bank. Lfidtflu 1741-)
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1 would like in express my gratitude to the followifig (07 15° 5°}? “"7 ha"
given me in tracing thcs: and other books: Dr 5-  Asdfll. M154 Ethtl G-
Bmdie, D: R. E. D. cm, Dr E. J. mngwau, Mm Elmer Gregory. Pr
Amald C. Klebs, Mr W. B. McDaniel, Dr Want! Pagcl and Mr H. Zur-
 
linger.


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Needham J. A History of Embryology. (1959) Cambridge University Press, London.

1959 Needham: Chapter 1 | Chapter 2 | Chapter 3 | Chapter 4

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A History of Embryology

Joseph Needham, F.R.S.

Fellow of Grmville E‘! Cain: College, and Sir William Dunn Reader in Biochemisty

in the University of Cambridge

Second Edition Revised with the Assistance of

Arthur Hughes, Ph.D

Lecturer in Anatomy in the University of Cambridge

Cambridge

At The University Press

1959 Published By The Synod Of The Cambridge University Press, London 0559: Benllty House


First edition 1934 Second edition 1959


Contents

Preliminary Note

List of Plates

Chapter One - Embryology in Antiquity

  1. Ideas of Primitive Peoples
  2. Egyptian Antiquity
  3. Artificial Incubation
  4. Indian Antiquity
  5. Hellenic Antiquity; the Pre-Socratic
  6. Hippocratic Embryology and the Doctrine of the Two Seeds
  7. Aristotle’s great Systematisation
  8. The Doctrine of the Menstrual Blood
  9. Denials of Maternity and Paternity
  10. Formation, Recapitulation and Fermentation
  11. The Aristotelian Balance—sheet
  12. Aristotle’s Theory of Causation
  13. The Hellenistic Age
  14. Galen and the Vital Faculties

Chapter Two - Embryology From Galen To The Renaissance

  1. Patristic Speculation
  2. Contributions of Jewish Thinkers
  3. Embryology among the Arabs
  4. Alchemy and Embryology
  5. The Visions of St Hildegard
  6. Albertus Magnus; the Re-awakening of Scientific Embryology
  7. Aristotle’s Masterpiece
  8. Scholastic Ideas on Generation
  9. The Insights of Leonardo da Vinci
  10. The Macro-Iconogrnphers of the Sixteenth Century
  11. The Movement to Rationalise Obstetrim

Chapter Three - Embryology In The Seventeenth Century

  1. The Opening Years
  2. Developmental Determinism and Trasplantation; Digby, Higlunom and Tagliacozzi
  3. Thomas Browne and the Beginnings of Chemical Embryology
  4. William Harvey and the Identification of the Blastoderm
  5. The Riddle of Fertilisation
  6. Harvey's Achievements and Influence
  7. Atomist Theories of Embryonic Development; Gaucndi and Dcscm-ta
  8. Fixatives and Uterine Milk; Robert Boyle and Walter Needhzun
  9. The Discovery of the Follicles of the Mammalian Ovary
  10. The Micro-iconographers and Preformatianism; Marcello Malpighi and Jan Swammerdam
  11. Foetal Respiration and Composition; John Mayow and Robert Boyle

Chapter Four - Embryology In The Eighteenth Century

  1. Theories of Foetal Nutrition
  2. Growth and Difierentiation; Stahl and Main-e-Jan
  3. Chemical and Quantitative Approaches to the Origin of Organisation; Boerhnave, Hamberger and Mazin
  4. Albrecht van Haller and the Rise of Techniques
  5. Embryos and Theologians
  6. Ovism and Animzdculism
  7. Spontaneous Generation
  8. Preformation and Epigenesis
  9. The Closing Years

Conclusion

Bibliography


Traditional methods of incubation

Portrait of William Harvey, act. 6t (1639)

Preliminary Note

The contribution to the history of science contained in the following four chapters first appeared as the opening part of a treatise on Chemical Embryology, published in 1931. They were delivered in the form of lectures about the same time at the University of London under the title “Speculation, Observation and Experiment as illustrated by the History of Embryology." The munificence of that University assured their appearance in separate, and amplified, form.‘

I suppose that the study of the history of science needs no apology. If at first sight the discussion of what was thought in the past rather than what is known now appmrs to be of merely antiquarian value, a deeper consideration will admit, with Louis Choulant, that the history of science is the guarantee of its freedom. The mistakes of our predecessors remind us that we may be mistaken; their wisdom prevents us from assuming that wisdom was born withus; and by studying the processes of their thought, we may hope to have a better understanding, and hence a better organisation, of our own. Theoreticil errors, such as the final cause, preformationisrn or phlog-iston; practical errors, such as the divorce between speculation and technique in the Hellenistic age, are always able to show us a more excellent way.

The present contribution does not claim, what probably no historical work can tnrly deserve, the ascription of a complete lack of bias in its presentation. Designed as it was to introduce a discussion of the border- line between embryology and biochemistry, it sought rather to lay bare the roots of chemical embryology in history, than to collect data indis- criminately on all the interesting aspects of the subject. Its title, “The Origins of Chemical Embryology," made no secret of this. And no obvious disadvantage attaches to such a plan, except the difficulty of deciding when to leave off. For although it is possible in reasonable space to try to dojustice to all aspects of embryology before 1800, after that date the number of investigators and thevariety of problems attacked becomes too great to handle conveniently on the same scale as before.’

‘ By embryology we mean in this book the embryology of animal: txduxivlly. The history ofthe embryology of plant: has been fully written only in Russian, by Bannov, but there is 1 shorter work by Souéges in French.

I or. the valuable work of Srudmflu; Florian; Dogelb; Oppenheimer; Fischer 5: Schopfer; and others.

Bifurcation bcgms; the spheres of morphology and ph,11o1ogy more obviously separate, and In the latter division chemical researches play an ever-increasing part. It is now hoped that a group of workers Wlll soon be able to continue the story in a companion volume through the nineteenth century under a number of separate hadings.

No exhaustive treatise on the history of embryology as yet exists.‘

The nearest approach to it is the very valuable memoir of E. Bloch with its epitome. but this only covers the era of the Renaissance with thor- oughness. Hertwig's account, which he printed at the beginning of his great Haruibuth tier Enlwicfalungxlehre, does not deal very fully with any aspect of the subject before x8oo, nor do the much shorter ones of Hen- neguy and Minot. The latter paper is interesting in that it ends with an emphasis on the need for physico-cherniczil work in the future. The introduction to Keibel's book is much slight;-r, but contains some useful information. There are various monographs and papers on special points, such as Youchet's rather untrustworthy treatment of the em- bryology of Aristotle, and Lones’ discussion of it, which is worse. Camus‘ notes are still the best commentary on the I1z'.rtlm‘a Animalium. Again, useful information on some cultural points is to be had from the treatise ofl"loss 6: Bartels. The introductions to certain books also con- tain valuable information, and in this class comes Dareste's remarkable book on temtology. The bibliographies contained in Von Hallefs eighth volume and in the books of Schurig and Hefiter are naturally of the greatest assistance. The valuable books of F. J. Cole and Thadeusz Bilikiewicz on seventeenth~century embryology appeared too late for use in the first prepamtion of this book, but have contributed to its revision.’

In 1939 there appeared a work, The Rise of Ermlryalagy, by the learned Californian anatomist A. W. Meyer, author of numerous periodical publimtions on our subject, some of which are referred to in the bibliography. His book stands to mine in much the same relation as the second volume of David Eugene Smith’s notable History qfMaihe- malice to the first; the one adopting 2| basically chronological treatment, the other A topical form in which separate subjects are chosen in succes- sion for consideration. However, Meyer devotes the bulk of his work to

I am we cannot attempt to provide a bibliography of the more upP°¢,m'fl modem work dealing with the subject melt". Yet in case suenufic mm or historians of other fields might appreciate some helpful introduction to embryology. Inlntton rnly ‘>9 made oi the popular boolu of Rouund, Wnddington mil Guttnnrher. An engineer or on hillflnan of xstronmny might lhlfl proceed to the recent tunes: of Wlddington,

man, or Willier 2: al. _ . - Certain minor works on the history of embryology hive proved maee¢sxble— Beulre; Eccleahymer; H. Fnheuder; Fnvaro; l’-‘enzkel; Gills: Hapr; Ottow. Other

urtides deserving mention are those of Gerber; Keller; du Ems.


the eighteenth and nineteenth centuries, passing over the earlier periods in his first thirty pages. His treatment of the nineteenth century is in- teresting indeed, though nothing could supersede the remarkable work of E. S. Russell, Form and Function. Particular interest attaches to L. A. Blacher’s monograph on Embryology in Russia in the 18!]: and 191/: Centun'e.r (195 5), since so much of the classical work centring around 1800 was done or published in that country.

These observations made, the principal reviews of the subject are chiefly to be found in histories of science in general, such as Sartou’s; histories of biological theory, such as R.’idl’s; histories of obstetrics, such as von SieboId's, Spencer's and E. F asbender's; histories ofgynaecology, such as McKay’s; and histories of anatomy, such as Singer's and Von T6ply's. Histories of medicine as a whole are numerous and good: I have found those of Garrison and Neuburgcr-Pagel most useful. Those which deal with special periods are also of assistance, such as Schrutz and Browne on Arabian, I. Bloch on Byzantine, and I-larnack on Patristic medicine. Histories of chemistry provide no help, for ancient chemistry was so oriented towards “practical” results, such as the lapi: philorophomm and elixir vitae, that the egg was only considered as a raw material for various preparations. The investigation of its change of properties during the development of the embryo did not occur to the alchemists. Detailed studies of particular subjects, such as those con- tained in Singer's two excellent volumes, The History and Method of Science, may also be of some assistance. Again, there are books which give a wonderful orientation and an articulate survey of vast tracts: of these Clifford Allbutt’s Greek Medicine in Rome, with its mass of refer- ences, is among the most valuable. And Miall’s Earbv Naturalist: must not be omitted, far, apart from the peculiar charm of style which marks it, it contains some singularly helpful bibliographical data.‘ But the study of the original sources, so far as that is possible, is a duty which cannot be avoided, and in what follows I have been careful to copy down no statement from a previous review when it was possible to read the actual words of the writer himself. This practice of going to the originals is made peculiarly necessary in a case such as the present one, when the history of a subject is regarded from a rather new angle.

The arrangement of my chapters I adopted in the first edition, and now preserve, only on the ground that it is suitable enough in the pres~ ent state of historical knowledge. Little was then said about eml>ry~ ology in China beuxusc at that time I could find out little about it, but it will be thoroughly treated in the eighth volume of my work on the

‘ A fine beginning has been made on the bibliographies of seventeenth-century mm of science by Keyna and Fulton.

history-of science in general in that great culture, Scimte and Ciril- "“"""_ W C{'1"0- Nor am I content with the short section on embry- ology In India, but here there are special difficulties owing to thc absmce of an established chronology for ancient and mediaeval Indian texts and an adequate account of it must be left for others to give. No per: manent framework for historical facts is proposed in what follows; I only attempt to bring them together, and to reveal some of the relation. ships between them. If the traditional pattern turns out to be badly distorted-—and there are many signs that it may—the facts can be rearranged.

But in whatever way this may tum out to be desirable, one neeasity rnust constantly be kept before the mind's eye, namely the knowledge of the relations between scientific thought and technical practice at any given period. For embryology this knowledge is diflicult to acquire, since up to the time of the Renaissance obstetrics remained a part of primitive folk-medicine rather than of serious medical science. We see, however, in the publication of the Hellenistic gynaecological treatises in the sixteenth century (Bauhin, Spach; see p. 109) the satisfaction of a new demand, even though it took the typiml Renaissance form of what might be mlled palaeolatry. It was part of that movement to rationalise obstetric: which included Harvey’s De Gmnatione and Malpighi's De Formatione Pulli and culminated in the celebrated man-midwives of the eighteenth century.‘ Again, the relation of the early systematists— Belon, Rondelet, Aldrovandus, Ray-—to the beginnings of mermntile expansion is fairly clear, for the mediaeval bstiary could not cope with the influx of new animals and plants from hitherto unknown regions, any one of which might prove to be an exploitable commodity.

The Hellenistic divorce between scientific thought and empirical technique is an important case in point. Greek life was divided strictly ima amela and ngdftg. The latter was not thought fitting [or a man of good birth. "Antiquity," says Diels. “was entirely aristocratic in attitude. Even prominent artists. Such 35 P5555335. W619 0135355 33 fifti- sans, and were incapable of bursting through the barrier separating the workers and pmsants from the upper clam. A second muse of the slight technical progress in antiquitywas its slave-holding system, which led to a lack of any impulse to develop the machine as a substitute for manual labour.” Xenophon in the Oemnomicus held the industries in poor repute.‘ "Men engaged in the mechanical arts,” he says, “must ever be

‘!3.g. the Chnmberlenx. Palfyn (see Stein), Mluricuu, _VVfl.ltarn Srnellrz, John Burton of York ("Dr Slop"), and Joseph Needhnm or Dewnzes; see the Imeles or Rosemhnl and Mengert. The dissertation ofCaspar Base (1729) is I typuml attack on the midwives ufllll time.

‘ See Creeotti. ' xv. 3; VI. lJ'l5-

both bad friends and feeble defenders of their country." He troubled himself little with those skilful in carpentry, metallurgy, painting and sculpture, but was always anxious to meet. a “gentleman" (ualzig rs miyafldg). The results of this were inevitable. Classical surgery and obstetrics benefited practically nothing from the speculations of the biologists from Alcmaeon to Herophilus (see pp. 29 Surgeons and midwives remained members of the painter-cobbler-builder group, the group of base-bom "rnechanicks”, entirely distinct from the astronomer- mathematician-xnetaphysician-biologist group, the group familiar with courts and tyrants.

Only the greatest broke away from this tradition: Aristotle, when he conversed with fishermen, Archimedes perhaps, when he constnicted his mechanical devices. For the rest, it was too strong. Down to the end of the Roman period the artillery in use remained precisely what it had been six hundred years before, although the Empire was crumbling under barbarian pressure, and would have given anything, one would imagine, for an improved artillery capable of withstanding the Gothic armies. It is strange, as has been acutely said, that the Rornans never invented anything so much in the Roman taste as a railway. So far as Hellenistic empirical industrial chemistry was concerned, the Demo- critcan and Epicurean atoms might never have existed. And in medicine, the only effect of the brilliant Greek atomic speculations was to give rise to the Methodic school of Roman physicians, described by Allbutt, whose influence was never strong, and who contributed relatively little to the main stream of therapeutics originating with Hippocrates.

In sum, we must not dissociate scientific advances from the technical needs and processes of the time, and the economic structure in which all are embedded. We shall never understand the failure of Greek science if we consider it in abstraction from the environment which sterilised its speculation. The history of science is not a mere succession of in- explicable geniuses, direct Promethean ambassadors to man from heaven. Whether a given fact would have got itself discovered by some other person than the historical discoverer had he not lived, it is certainly profitless and probably meaningless to enquire. But scientific men do not live in a vacuum; on the contrary, the directions of their interest are ever conditioned by the structure of the world they live in. Further historical research will enable us to take into account the social and economic status of the invatigator himself (cf. Chambers for the Hellenistic artist, and Yearsley for the sixteenth-century physician).

It would thus be of the greatest interest to lmow accurately the sources of the emoluments of embryologists at different times.‘ From Om-

‘ On this, cf. Cumston and Dittrielr.

steinjstadmirzible book on the scientific societies of the Renaissance, the suspicion arises that their royal patronage was dictated not only by

1 purely disinterested passion {or abstract truth, but by a desire to profit as much as possible by the new techniques which the decay of the gnfi- usury doctrines, the Willingness of the rising mercantile class m mkc mdusmfil "ffimlllts." and um f”'1'3“8i"8 thought of the scientific men were combining to produce. In England's Royal Society, indeed, me preoccupation of the early Fellows with the Uirnprovexnent of u-ad; and husbandry” is patent to anyone acquainted with its early history (cf. Thomas Spint’s account of it)! Thus Dr Jasper Needham, clccled in 1663, read only one paper before the Society—not, as might haw; been expected from his profession, on the transfusion of blood or the anatomy of the brain; but on the value and use of “China Varnish". However, it is probable that for the most part the emhryologists whose work we shall have to discuss were pi-.iciisi.ng physicians, free or relatively free from the ancient tnidition, and conscious that to understand the mystery of generation would be to advance the science and art of medicine.

In this connection it is of interest that the Church in the seventeenth and eighteenth centuries provided a certain source of demand for em- bryological research. Of this Swaminerdarn and Malehranche (see p. x69) provide interesting examples, and the conviction, then widely held, that research into the nature of generation would throw light an orthodox theological doctrines, such as that of original sin, led to an economic situation of value for biological development. Finally, it would he rash to minimise the factor of pure curiosity in seventeenth- century science. The recreational quality of Lecuwenhoek's investiga- tions is, as Baas-Becking says, too obvious to be overlooked.‘

The history of single forms of scientific knowledge is in way hap- pier because containing more of continuity than that of civilisation as a whole. The assiduity with which men of diflerent periods in the rise and decline of a culture pursue the diflerent forms of human experience may, as Spengler has shown, vary much, but those forms remain funda- mentally the same, even if their manifestations are profoundly changed.


scphy, the mechanic, and husbandry, according to the principles of our new philoso- phical college, that Values no knowledge, but as it hath a tendency in use.And therefore I shall make K on: of my xuiu to you, that you would take the pains to enquire lllILl€ more thoroughly inro the way: of husbandry etc. practised in your pan}: ‘rid when you intend for England, to bring along with you what good receipt: or choice books of my of those subjects you can procure; which will make you extremely welcome to our

‘ ‘b I , hichlhad d ' edt ‘ oulil 'ti of.”Fultonre- imzmmi of nuieiiifé but wisgiapy taaliie leave to think

it was not to inadequate as um-rywould suppose. _ . - The run loop: of Leeuwenhoek‘: discoveries Is now appearing, drum to the labour: of van Runberlc and his mllabontvfi.


That science, at any rate, does maintain some sort of continuity what- ever gaps there may be between the phases of its progress, is a belief agreeable with all the available facts, and one which no criticism will easily shake.

It only remains to record my indebtedness to those who have assisted me in the preparation of this work. Primarily I am grateful to Dr Charles Singer, who annotated my typescript with valuable comments and lent me many papers and pictures, and to Professor R. C. Punnett, who placed unreservedly at my disposal his knowledge of the history of generation and his library of old and rare biological books. To Dr Arthur Peck I am indebted for the correction of my Greek, and it was Professor A. B. Cook who introduced me to the embryology of the ancients. For guidance on Talmudic and Jewish matters I thank Dr Walter Pagel, the late Dr Louis Rapkine and Dr H. Loewe. Without the assiduous backing of Mr Powell, the Librarian of the Royal Society of Medicine, and his assistants, and of Mr H. Zeitlinger, I should have dealt much more inadequately than I have with papers and books which cannot be consulted in Cambridge. And in addition to those mentioned above, the following friends kindly read through and criticised the proofs: Pro- fessor Reuben Levy, the late Professor F. M. Cornford, the late Sir William Dampier, Mr Gregory Bateson, Professor Roy Pascal and the Rev. W. L. Elmslie.


To the Master of Gonville and Caius College I am indebted for permission to reproduce the portrait of William Harvey (attributed to Rembrandt) which hangs in our Senior Combination Room. Although the authenticity of this is not accepted by Keynes in his recent study of the portraits of Harvey, it has been in the possession of the College since r798, when it came to us from the Earl of Leicester. After com- parison with other portraits of Harvey, many feel unable to concur in its rejection.


Plates

(A) Egyptian peasant incubator (from Cadman) (B) Chinese peasant incubator (from King)

Heteromorphosis in a guardian deity (lokapfila) depicted in a fresco on the wall of one of the m‘vc~tempIes at Ch’ien-F0-Tung, Tunlnmng, Kansu province, China

The oldest known drawing of the uterus. From a ninth- century MS. (the Brussels Mosdlion Codex) of Soranus’ work on gynaecology

An illustration from the Liber Sn'1:z'as (LAD. 1150) of St Hildegard of Bingen (Wicsbaden Codex B), showing the descent of the soul into the embryo (after Singer)

A page from Leonardo da Vinci’s anatomical notebooks (guaderni d’Anazmm'a), c. AD. 1490

Portrait of Volcher Coiter, act. 41 (painted in r 575 byan unknown master)

Portrait of Sir Kenelm Digby (from the painting by Oomelius Jansen, ca. 1650)

Portrait of Sir Thomas Browne and his wife, Dorothy (ca. x650)

IX Zeus liberating living beings from an egg (the frontispieoe

of William Harvey’s book on the Generation of Animal:, 1 6 51) ‘

X Illustrations from “falter Needl1am’s De Fonnatn Foetu of 1667

PLATES

Illustrations from Maipighfs 1): 01:0 Incubate of 167: shgwmg the early stages of the development of the °h“'-k facing page 166

Illustrations from 77:: auiou: and accurate ab.mvan'an: 0/ Mr Stephen Lorene-t'm' of Florence on the Dirxrrtioru qf

the Cramp-Fish ,9, Metamorphosis in Buddhist iconography; statues in the

Sleeping Buddha Temple at Suchaw (Chin-dffian),

Kansu province. China ,7; Portrait at‘ Antan van Leeuwmhoek by Job.

Vex-kalje (1686) :7: Portrait of Robert Boyle (a. 1690) 176 A tel-atoms with well-formed teeth and hair (from Robert

Plat‘: Natural Hi:-tmy afS!ufl'ord:h:'r:, 1686) 178 Illustrations from Antoine Maine-Jan‘: Obaematiovu

mr Infmnahbn du pcmlet (1722) 186 De Réaumux-‘I Incubators (from De (‘art de faire (chm

In paulm, 1749) 204

Note - The use of the Ihonened and (&) indium eollzbcmion between two or more author


Cite this page: Hill, M.A. (2024, April 27) Embryology Book - A History of Embryology 1959. Retrieved from https://embryology.med.unsw.edu.au/embryology/index.php/Book_-_A_History_of_Embryology_1959

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