Book - History of embryology in Russia 1750 - 1850: Difference between revisions

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==Contents==  
==Contents==  


Introduction by Jane Maienschein
[[#Introduction by Jane Maienschein|Introduction by Jane Maienschein]]


Preface
[[#Preface|Preface]]


# [[Book - Russian Embryology (1750 - 1850) 1|The Beginning of Embryological Investigations in Russia in Lomonosov's Epoch]]
# [[Book - Russian Embryology (1750 - 1850) 1|The Beginning of Embryological Investigations in Russia in Lomonosov's Epoch]]
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# [[Book - Russian Embryology (1750 - 1850) 25|Investigations on Invertebrate Embryology - Work of A. Grube, A. D. Nordmann, N. A. Warnek, and A. Krohn]]
# [[Book - Russian Embryology (1750 - 1850) 25|Investigations on Invertebrate Embryology - Work of A. Grube, A. D. Nordmann, N. A. Warnek, and A. Krohn]]


Comments
 
[[#Comments|Comments]]


Illustrations To Blyakher, History Of Embryology In Russia
Illustrations To Blyakher, History Of Embryology In Russia


==General==
==Introduction by Jane Maienschein==
 
===General===


When Dr. Robert Multhauf asked me if I would consider editing this translation of Blyakher's volume, he warned that this was part of what seemed to him a most unusual scholarly project. Thanks to a somewhat mysterious and complicated government exchange program, the Smithsonian Institution and the National Science Foundation had been charged with overseeing the translation into English of several foreign language texts in the history of science. Upon the recommendation of experts, the volumes chosen included two by L. Blyakher, a Russian biologist. In particular, these Russian volumes, including THE HISTORY OF THE INHERITANCE OF ACQUIRED CHARACTERISTICS, edited by Frederick Churchill, and this one, were thought to present a valuable exposure to a Russian point of view in the history of science and to detail important episodes of Russian scientific history. Therefore, the translation began.
When Dr. Robert Multhauf asked me if I would consider editing this translation of Blyakher's volume, he warned that this was part of what seemed to him a most unusual scholarly project. Thanks to a somewhat mysterious and complicated government exchange program, the Smithsonian Institution and the National Science Foundation had been charged with overseeing the translation into English of several foreign language texts in the history of science. Upon the recommendation of experts, the volumes chosen included two by L. Blyakher, a Russian biologist. In particular, these Russian volumes, including THE HISTORY OF THE INHERITANCE OF ACQUIRED CHARACTERISTICS, edited by Frederick Churchill, and this one, were thought to present a valuable exposure to a Russian point of view in the history of science and to detail important episodes of Russian scientific history. Therefore, the translation began.
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July 1981 Arizona State University
July 1981 Arizona State University


==Preface==
==Preface==
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The author is deeply grateful to T. D. Detlaf; L. D. Lioznera, S. R. Mikulinsky, S. L. Sobol and G. A. Shmidt, who listened to the reports about the contents of this book or read the manuscript and made critical comments. The author particularly thanks the collective of the library of the Moscow Society of Naturalists, who willingly helped during the search for literary sources and illustrated materials.
The author is deeply grateful to T. D. Detlaf; L. D. Lioznera, S. R. Mikulinsky, S. L. Sobol and G. A. Shmidt, who listened to the reports about the contents of this book or read the manuscript and made critical comments. The author particularly thanks the collective of the library of the Moscow Society of Naturalists, who willingly helped during the search for literary sources and illustrated materials.


==Chapter 25. Investigations on Invertebrate Embryology - Work of A. Grube, A. D. Nordmann, N. A. Warnek, and A. Krohn==
During the first twenty to thirty years of the nineteenth
century, embryology remained chiefly the study of the embryonic
development of vertebrates; the comparative peculiarities
of development of different animals was studied only at the
limits of this most studied group. By the end of the twentieth
year of that century, i.e. the period of Baer's active work
in embryology, the first investigations into the development
of invertebrates appeared. Baer himself, as mentioned above, 1
turned his attention to the characteristic peculiarities of
development of arthropods, noting in particular that their
blastoderms are situated on the abdominal side of the egg and
distributed from here in the dorsal direction. Baer was not
able to explain the development of arthropods more clearly.
Even the investigations of his predecessor Herold, which were
limited to the study of the late stages of development of
butterflies and spiders, 2 could not give Baer material for
well-grounded comparative embryological conclusions.
The aspiration to apply embryological principles to
arthropod development was actualized by Baer, first in the
study of embryonic layers, reflected in the works of
1. See Chapter 15.
2. J. m. D. Herold, ENTWICKELUNGSGESCHICHTE DER SCHMETTERLINGE ANATOMISCH UND PHYSIOLOGISCH BEARBEITET
(Cassel u. Marburg, 1815), vi + 118 + xxxiv pp.; EXERCITATIONES DE ANIMALIUM VERTEBRIS CARENTIUM IN 0V0
FORMATIONE. DE GENERATIONE ARANEORUM IN 0V0
(Marburg, 1824) , x + 63 pp.
M. H. Rathke. A native of Danzig, Rathke in 1829 arrived
in Russia, where he was professor for six years at Dorpat
University. Before that, he published many embryo logical works,
including a valuable work on the development of crayfish. 3
During his tenure at Dorpat, Rathke visited Moscow and
Petersburg and also travelled in the Crimea to investigate
the fauna of the Black Sea. On the Black Sea coastline,
Rathke collected comparative embryological material which
was later used in ON MORPHOLOGY: TRAVEL NOTES FROM
TAURIA.4 in this collection, in addition to brief information
about the embryonic stage of actinia, there is also an essay
on the embryology of the Crimean scorpion and investigations
concerning the development of nine species of Crustacea of
different orders (copepods, araphipods, decapods, and isopods) .
Rathke 1 s work on crayfish development, and also his investigations on the development of other arthropods (124), represent
a clear interest in describing the phenomenon of embryonic
development in arthropods according to the ideas of Pander and
Baer. Rathke spoke of the embryonic disk or blastoderm, of
the primary cavity, and of the two embryonic membranes (serous
and mucous) into which the blastoderm is divided. The first
stages of development, the division of the ovum and the first
processes of separation of the rudiments, remained untraced.
It must be borne in mind that a clear presentation about
the essence of the processes which take place in the early
stages of embryonic development — i.e. first of all the process
of division — was not yet established in the first forty years
of the last century. Thus, Reichert (125) studied the development of the frog ovum but reached an incorrect conclusion
about the structure of the still-undivided ovum, supposing
that it consisted of many cells (by cells he meant the round
accumulations of the yolk plates) . This point of view was
raised by Reichert both in his DIE ENTWICKELUNGSLEBEN
IM WIRBELTHIRREICH and in an article published one year
before, "On the Process of Division in the Ova of Amphibia, "5
3. H. Rathke, UBER DIE BILDUNG UND ENTWICKELUNG DES
FLUSSKREBSES (Leipzig, 1829) , 97 pp.
4. Rathke, ZUR MORPHOLOGIE. REISEBEMERKUNGEN AUS TAURIA
(Riga und Leipzig, 1827) , 192 pp.
5. K. B. Reichert, DIE ENTWICKELUNGSLEBEN IM WIRBELTHIERREICH (Berlin, 1840), x + 261 pp.? "Uber den Furchungsprozes
der Batrachier," ARCH. ANAT . PHYSIOL. (1841), pp. 523 541.
514
in which he wrote: The process of division of the amphibian
ova is nothing more than successively accomplished generic
action (GEBURTSACT) of the maternal cells, repeatedly
invested in each other.
T. Bischoff, working on the embryology of mammals and
publishing in the period from 1842 to 1852 a monograph
about the development of man, rabbit, dog and guinea pig,
also did not reach a clear understanding of processes of
division and did not recognize the spheres resulting from
division as cells; since in his opinion, the cell must
possess a cavity, but the spheres resulting from division
are filled with yolk, the nuclei of the blastoderms were
taken by Bischoff as fat droplets.
A. K8 Hiker** went much further in the analysis of the
process of ovum division, admitting the direct continuity
of the blastoderms and those cells from which the embryo is
built in later stages.
In all these cases the discussion was about the forms of
development, complicated by the great quantity of yolk in
the centrolecithal and telolecithal ova. The nature of the
occurrence there of superficial and discoidal division was
explained much later, after the introduction of sectioning
in embryology.
A more distinct presentation on the phenomena of
division was stated by Baer in his work on the development
of amphibia, 7 but especially in the work noticed by his
contemporaries and later forgotten on the development of the
ova of the sea urchin. 8
Certain embryologists of the thirties and the forties
nearly approached the correct interpretation of the phenomena
of the ovum division. They include KClliker, then Loven,9
6. A. KSlliker, ENTWICKELUNGSGESCHICHTE DER CEPHALOPODEN (Zurich, 1844), 180 pp.
7. See Chapter 21 .
8. See Chapter 23.
9. S. L. Loven, "Bidrag til kannedomen af Molluskenas
untveckling," K. VET. AKAD . HANDLINGAR (1839),
pp. 227 - 241.
Sars,10 van Beneden,H and Quatrefages-^ and must be
mentioned. The first four investigated mollusc development,
and the last studied the development of annelids. They
presented some stages of ovum division, but did not broach
the subject of the internal processes occurring in it nor
of the fate of the spheres of division.
A great part of the works of that time concerning the
development of invertebrates was illustrated by the study of
different types of reproduction, and also by the description
of the structure and the transformation of different larval
forms which sometimes did not yield to systematic determination and figured under different specific names (126) . The
investigations of forms of reproduction became especially
popular after Steenstrup showed the wide distribution of
the alternation of sexual and asexual generations of many
invertebrates; the application of this empirical regularity
to groups of animals not investigated before this time
constituted the majority of works in the present sphere.
The single base for generalization concerning early embryonic
development is thought to be the cellular theory formed
shortly before this. Using extremely imperfect microscopic
techniques, the embryologists of the first half of the
nineteenth century posed the question, can the spheres of
division be called cells, and are the cells of which the
embryo consists the direct descendants of the primary
spheres of division? They attempted to trace the fate of
those existing in the unfertilized ova "embryonic vesicle"
(and "embryonic speck"), i.e. to explain whether these formations disappear after fertilization without a trace or
whether they stand in continuous genetic connection with
the nucleus (and the nucleolus) of the cells of the embryo.
On the foundation of a one-sided and primitive understanding
of the cellular theory, fantastic presentations sometimes
grew, like the ideas of Reichert, which resurrected the
long-buried theory of preformation.
10. M. Sars, "Beitrag zur Entwickelungsgeschichte der
Mollusken und Zoophy ten," ARCH. NATURG . , 6 (1840),
pp. 196 - 219.
11. P. J. van Beneden et Ch. Windischmann, "Recherches sur
l'embryogenie des Limaces," ARCH. ANAT . PHYSIOL.
(1841) , pp. 176 - 195.
12. A. de Quatrefages, "Sur l'embryogenie des Annelides,"
ANN. SC. NAT., 3 Ser. Zool. 8 (1847), pp. 99 - 102.
For the purposeful coordination of the efforts of
zoologists studying invertebrate development, the theory of
embryonic layers, which still had not become a broad
scientific generalization, could not serve. Its correctness
was proven only for vertebrates, but the application of
the theory of embryonic layers to invertebrates was believed
by nearly no one. Rathke's old data concerning crayfish were
known. More than a quarter of a century later Zaddach also
reported on the embryonic layers of insects, admitting to
his descriptions crude morphological mistakes.
The data related to the development of different types
of invertebrates were accumulated relatively slowly, because
a theoretical conception for which this material could be
used did not exist. The study of the types of structure and
development of animals, taking its beginning from the opinion
of Cuvier and Baer, fell into decay. The formally opposite
theory of types, that of Naturphilosophie, the idea of
unity of planes, caused little enthusiasm. Only after fifty
years, after the appearance of Darwin's theory, did this
period of mental stagnation end. Then the naturalists were
divided into two camps — the hearty supporters and the violent
opponents of the theory of evolution. Practically none of the
contemporaries of the great reformer of biological science
could keep an olympian calm and sustain neutrality for any
length of time.
The investigations preceding the publications of the
ORIGIN OF SPECIES played, however, its historical role.
Factual material was collected which Darwin and his early
followers used. In this period came the important works of
the Russian embryologists A. Grube, A. D. Nordmann,
N. A. Warnek, and A. Krohn.
To Dorpat University professor A. Grube belonged serious
investigation on the development of annelids; he studied
the embryonic and partly post -embryonic development of the
Proboscidea leeches Clepsine oomplanata and C. biooulata
(genus Glossosiphonia, according to the current terminology). 13
Adolf Edward Grube* 4 was born in 1812 in Konigsberg,
where he graduated from the university. For thirteen years
C1843 - 1865) he was professor of zoology and comparative
anatomy in Dorpat University, and to this period of Grube's
life are related his most important works in systematics,
anatomy, and embryology of the annelids.
Grube's work undoubtedly possesses remarkable significance. Because it was undeservedly forgotten and not
mentioned even in the detailed reports and special works on
the development of leeches (127), we must dwell here on its
content.
In the introduction to his work, Grube turned to
memories of his years of study in Konigsberg, where Baer
was working. Actually the great embryologist at that time
read few lectures, being engrossed in the investigations
of development of fish and amphibia. He was helped by
Grube's friend, the clever graphic artist Burov, who, by
the way, encouraged Grube's attraction to the laboratory,
where the latter acquired a taste for embryological
investigations. Grube was aware of the difficulty of these
observations, but he did not abandon his idea of devoting
himself to embryology.
13. A. E. Grube, UNTERSUCHUNGEN UBER DIE ENTWICKLUNG
DER ANNELIDEN. I: UNTERSUCHUNGEN UBER DIE
ENTWICKLUNG DER CLEPSINEN (Konigsberg, 1844),
ii + 56 pp.
14. The author expresses heartfelt gratitude to the head
of the Department of Zoology at the University of
Tartu, Professor Kh. Kh. Riikoya, for the photo
reprinted here of A. E. Grube.
Shortly after graduation from the university, Grube
left for a trip to the Mediterranean seacoast. "Baer's
observations on the history of animal development," Grube
wrote, "accompanied me on the trip to Italy, and along with
the enjoyments of nature and art I continuously sated my
interest in this sphere of science" Cp. 1). On the coast
Grube zealously collected zoological materials and made
dissections, considering these studies important for his
future embryological investigations. His attention was
attracted mainly to annelids, the study of which became
his basic zoological specialty. Only in 1839 did Grube turn
to work directly on annelid development, beginning with
the study of the very small ova of the Saenuris vaviegata
described by Hofmeister. In addition he reported that their
development differs in many relations from the corresponding
phenomena in medical leeches which had been very superficially
described by E. Weber. The following spring, Grube studied
the embryology of the leech and confirmed the reality of
the formerly discovered differences. Preparing the results
of his observations for publication, Grube conscientiously
studied the works of the authors whom he considered his
predecessors. The work of Filippi on the anatomy and
embryology of Proboscidae leeches,! 5 recently published,
especially interested him. Grube found that in Filippi' s
investigations there was only scanty information about the
division of the ovum, as the Italian author saw only "six
lobes, situated in one and the same plane around the seventh,
situated in the center." To this description he added that
these spherical segments disintegrate into smaller spheres
(GLOBULI ORGANICI), sharing in the formation of the
embryo. Filippi apparently studied the developing ova through
the membrane of a cocoon, which can explain the indistinct
results of his observations. The first foundation of the
embryo Filippi described with an indefinite expression-— cutis
(LA CUTE) . He also erroneously considered, that the
hatched young is nearly similar to the adult leech and at
once is attached to the mother by the help of posterior
suckers (in fact they are not present at this time) .
Grube could trace the development of Clepsine in
incomparably more detail; his success was aided by the
application of concentrated reagents, in particular diluted
nitric acid. In the first section of his work, Grube
described the structure of the female and male sexual organs, copulation, the act of deposition of eggs, and also
the structure of the ova at the time of maturation and
directly after deposition.
15. F. de Filippi, LETTERA AL S. DOTT . RUSCONI SOPRA
L'ANATOMIA E LO SVILUPPO DELLE CLEPSINE (Pavia,
1839) .
The still-undeposited egg, hung in a special pocket
in the oviduct, consists of a fine-grained substance
(molecular bodies) and yolk (fatty bodies) ; it is provided
with a nucleus (embryonic vesicle) . Before the deposition,
the ovum, which is separated from the oviduct and is freely
situated in the egg-reservoir, has the same structure; the
nucleus already cannot be seen in a beam of light and can
be seen only during the crushing of the ovum. Later the
embryonic vesicle disappears completely.
The second section of the work was devoted to the
description of embryonic development. In the deposited ovum
for one hour no change can be seen, and then at its poles
the following phenomena can be observed. At the beginning,
on one of them, a white spot appears; this increases and
turns into a disk with a grey spot in the center (Figure 36, a).
Then this grey center increases and a white spot appears in
it, and the external white disk is transformed into a ring,
which Grube called the polar ring (Figure 36, b) . The field
of formation of the polar ring he called the active pole,
since according to his observations, it is here at the time
of development of the embryo that the most noticeable changes
take place. On the opposite inactive pole a white ring appears,
but less distinctly delimited. The polar ring ascends over
the surface of the ovum in the form of a papilla, so that
after its infiltration its condensation can be prepared.
Grube considered that the formation of polar rings is the
result of the displacement of an internal substance, during
which the "molecular bodies" are gathered in the form of
rings on the poles.
Following the appearance of polar rings, or, in Grube* s
expression, "the process of formation of fissures
(DURCHFURCHUNG) or, rather, cleft (ZERKLUFTUNG) . . . .
those fissures not only occur on the surface, but penetrate
deep into the mass of the yolk, as a simple experiment shows:
under careful pressure of the condensed yolk, it disintegrates
into as many parts as the segments are delimited by the
fissures" (p. 17). The first fissure divides the ovum into two unequal parts [Figure 36, c), so that the polar ring can
be divided into halves, or from it a small part is separated,
or the fissure occurs in direct proximity from the ring,
not touching it completely. The second fissure divides the
small segment approximately at a right angle to the first
(Figure 36, d), and the third also divides the large segment
(Figure 36, e] .
Thus, the ovum passes into the stage of four blastomeres
(Grube noted that the large segment in many cases is divided
earlier than the smaller one) . Of the four segments formed,
one is larger than all the others, and it is divided by a
fourth fissure which provides the beginning of the fifth
segment. The following division is concerned with this last
and takes place not along its length, but transversely, the
result of which is that the divided fifth segment forms a
region in the form of a polar field in the inactive pole
of the ovum (Figure 36, f) . The sixth, seventh and eighth
fissures again go from the active pole to the polar field of
the inactive pole. After the sixth division the polar ring
on the inactive pole usually disappears. Grube did not see
more than eight meridional divisions, and said that in
leeches the blackberry stage characteristic for many animals
is absent; in this stage the meridional divisions are replaced
by transverse ones.
At the same time as the division, accomplished by
means of fissures, in the ovum of Clepsine the process of
separation of the small globules from the large segments of
the ovum first described by Grube takes place; these globules
Grube called WANDUNGSBALLEN . In his opinion, the wall
of the body of the embryo is built from them; "Segments of
the yolk" and Wandungsballen are nothing more than the
macromeres and micromeres of the terminology of today's
embryology. The first small globule, as Grube noted, appears
on the active pole after the formation of the first fissure
and appears to be situated in this fissure. Judging by time
and place of the appearance of this globule, here the
discussion does not concern the formation of the micromeres,
but the separation of the first polar body. The following
small globules, forming on the active pole of the ovum,
undoubtedly are micromeres. Grube described their accumulation, ascertaining during this that by the increase of the
number of fissures the number of micromeres increases also;
however, these phenomena do not stand in an indissoluble connection (Figure 36, e and g) . The formation of micromeres,
according to Grube's description, is accompanied by the
replacement of substances inside the ovum segments; he
described these replacements in detail, alternating the
statement of facts about his observations with theoretical
conjecture about the forces of attraction, points of their
application, and so on.
The micromeres, accumulated on the active pole, do not
exhibit equal size; the smaller they are, there are relatively
larger molecular substances (protoplasm) anc i smaller fatty
globules (yolk) in them. The smallest consist only of protoplasm and a round transparent nuclear body. These micromeres,
according to Grube, never arise from macromeres; therefore,
he wrote, "I must conclude that the small globules originate
from those large globules already present on the pole which
earlier separated from segments of yolk" (p. 22). The
accumulation of micromeres forms a plate in the form of an
isosceles triangle, without sharp limits passing into the
other surface of the ovum. This plate, which Grube called
the embryonic field, is so situated that the summit of the
triangle is turned toward the active pole and corresponds
to the cephalic part of the future embryo.
It consists of "mosaic pieces" which arise as a result
of the multiplication of micromeres; in addition, these
pieces are smallest when they are nearest to the cephalic
end. In the stage described, not only the anterior and
posterior ends of the embryo, but also its upper and lower
sides, as the surface of the embryo, on which the embryonic
fluid is situated, correspond to its abdominal side. Using
the descriptive phrase "mosaic pieces", Grube resolved
consciously to avoid using the term cell, not being sure
that it inherits its own features, which are characteristic
of typical, in particular plant, cells.
The anterior end of the embryonic field quickly expands
(Figure 36, c) , and by forming two summits it produces S-shaped
curved shafts (Figure 36, j), which Grube called the abdominal
shafts. This name is well-founded, because these shafts,
appearing in the field of the active hemisphere of the ovum,
i.e. the future dorsal side of the embryo, are displaced
towards its ventral side, where they later accrete into a single ventral embryonic region. The ventral shafts,
according to Grube, on the ventral side of the leech embryo
play the same role which the dorsal shafts play in the
vertebrate embryos, because from them the wall of the body,
in particular its muscles, is formed. The ventral shafts are
composed of extremely small, closely adjacent other globules.
At the posterior end of the embryonic field remain large
globules which form part of its molecular (protoplasmic)
content; three of these terminal globules usually are present
on each side (Figure 36, k) . The internal part of the yolk
globule (the divided ovum) constitutes a mass containing
little protoplasm. This rich yolk mass is used mainly for
building organs which are present in the abdominal cavity,
in particular the intestinal canal; above it a layer grows,
composed from a molecular mass which takes its origin from
the ventral shafts.
Figure 36. The development of Clepsine (by Grube) .
a — fertilized ovum, b— ovum on which the white disk with grey
ring appeared, c— first fissure; the white disk is present
near it. d — yolk globule, divided into four parts, e— yolk
globule, divided by six transverse fissures; the polar field
increases as the surrounded globular segments decrease in size
due to the loss of albumen (molecular) mass used for the formation of WANDUNGSBALLEN on the active pole, f — -the same stage,
globules of division from the lower side; on the inactive pole
the polar field is seen, i.e. the surface of the seventh segment,
separated from the other round fissures, g — somewhat later
stage, view from, above; the number of WANDUNGSBALLEN increasing, they then form a small disk which can be regarded as a
rudiment (KEIM). h — WANDUNGSBALLEN increase in number and
occupy a great area, forming the embryonic field, i — the view
of the abdominal shaft from behind; they move so far apart that
they envelop the yolk globule by a ring . j— the abdominal shaft
from the side; on the posterior end of each, three whitish
globules are present, k — the abdominal shafts unite together.
The rudiment of the neural cord appears in the form
of two white stripes, joined to the external side by the
recently formed embryonic stripes. After their closure on
the ventral sides, the halves of the paired rudiment of the
nervous system are united.
In this period the surface of the embryonic body is
already covered with epithelium, composed of flat cells of
different sizes and forms. At the same time in the body
cavity dissepiments are situated, their number gradually
increasing.
In the third part of Grube f s work he included changes
occurring after the hatching of Clepsine from the egg
membrane. Contrary to Filippi, Grube found very essential
differences between the just-hatched leech and the formed
worms. In the short, cylindrical little worm hatching from
the egg, there are still no posterior sucking discs, no eyes,
no blood vessels, and the formation of the dissepiments is
also unfinished. As the young larvae are also immediately
fixed by the anterior end, Grube assumed the presence of the
rudiment of the anterior sucker with longitudinal and circular
muscles. Within one day after hatching this sucker is clearly
noticeable in the form of a convex ring; after two days
this ring elongates, as Grube thought, under the effect of the
heaviness of the body of the leech hanging on it. Inside the
elongated ring a canal appears representing, according to Grube, the rudiment of the sheath of the still unformed
proboscis. The wall of the intestinal canal, or yolk sac,
consists at this time of large cells. Later, by means of a
circular twist of the most anterior part of the body, the
caudal disc develops and the proboscis is formed, which the
larva can let out and pull in. The digestive canal is formed
in this way. On the intestinal canal extensions appear, and
in addition to this, it is divided into three parts. The
anterior and posterior parts are significantly thinner than
the middle one which composes the future stomach. In the last
the more enlarged lateral pouches appear. Later on the eyes
develop; at first they have the form of circular red spots.
The vascular system develops, and, on the dorsal side of the
body, the pulsating heart first becomes noticeable.
Grube finished the description of his observations with
the following words: "The development considered here is
probably spread extensively in the class ANNULATA. This
is established on the basis of detailed investigations on many
representatives of Naidae and Lumbricinae namely in Saenuris
variegata Hoffm., Euaxes acutirostris Gr. and Lwnbvioulus
variegatus Gr., and if it can be judged by an analogy with
the adult animals, it will be also correct for the genus
Lumbricus, and also for many sea worms" Cp. 45). With this,
Grube carefully observed that discussion of analogy without
special investigations can lead to erroneous conclusions.
A discussion of Grube' s work leads to the conclusion
that he promoted the study of embryology of the annulated
worms (annelids) and discovered phenomena the detailed study
of which was done significantly later. His most important
achievement in terms of recent embryology can be summarized
in the following way:
1. Grube established that in the eggs of Proboscidea
leeches there are polar plasmas; especially conspicuous are
protoplasmic rings on the animal pole.
2. He recognized the complete, unequal division of the
ovum of Proboscidea leeches, during which the animal polar
plasma can go in one of the blastomeres of the four-celled
stage.
3. Grube clearly saw how in the stage of the four
blastomeres, from the animal side, very small globules of
division began to separate; i.e. he established the fact
of the formation of micromeres, which are composed mainly
of protoplasm, and macromeres, which are rich in yolk.
4. Later he established that the number of micromeres
increased either by separation from macromeres, or by
means of multiplication of the early formed micromeres.
5. By Grube' s observations, the "rudiment" which is
formed on the animal pole, owing to the multiplication of
micromeres, spreads over the surface of the ovum, as a
result of which the macromeres appear inside the embryo.
This phenomenon very nearly resembles epibolic gastrulation.
6. At the end, Grube described the embryonic stripes
coming out from three pairs of terminal cells; i.e. he
discovered in the annulated worms the phenomenon of
teloblastic development.
7. The embryonic stripes, in Grube' s observations,
are displaced towards each other and are united on the
ventral side. From the material of the embryonic stripes
("the abdominal shafts") the wall of the worm's bodytegmens, muscles, and nervous system is formed.
Grube 's excellent work was forgotten, and superiority
in the study of the embryology of annulated worms, in
particular the leeches, was accorded to Rathke, whose work
was published eighteen years later. Rathke 's work without
doubt has merit, but also has many defects, so that it
can be seen as a step backwards in comparison with the work
of Grube. Thus in the maxillary leech Nephelis vulgaris
Rathke clearly saw the formation of micromeres, which in
Clepsine complanata he did not see, and instead of the
micromeres only granularity is illustrated in his drawing
in the animal parts of the blastomeres. At a later stage it
is shown as if the unlacing of the micromeres has begun, but
he did not see whether this led them to division. Together
with this, Rathke reproached Grube for seeking the source
of micromeres CWANDUNGSBALLEN) in the depths of the
globules of the division, while Rathke himself in general did not see their separation. According to his description,
a number of hillocks on the animal pole of the divided ovum
directly turned into a thickening which is the rudiment of
the embryonic stripes. The embryonic stripes of Nephelzs
are illustrated very schematically by Rathke; he did not
observe the characteristic cellular rows in these stripes,
neither did he recognize the connection between the only
three teloblasts illustrated in his drawing which are
situated as if they lie at the extreme ends opposite to
the embryonic stripes and the middle one as if it lies
between these ends. Rathke also represented the embryonic
stripes of Clepsine less distinctly than Grube.
All this is mentioned not to underestimate the
significance of Rathke ! s investigations, but only to drawattention to the more perfect work, in many aspects, of
Grube, to call to mind its priority and to show the importance
of his work in the history of Russian and world embryology.
In the 1830' s and 1840' s, the development of invertebrates also interested A. D. Nordmann (1803 - 1866). Aleksandr
Davidovich Nordmann was the son of a Russian officer born
in Finland. In 1821 he entered the university in Abo, and
after graduation he worked for some years in Berlin with
Rudolphi and Ehrenberg. During his stay in Germany he
accompanied Oken, Tiedemann, and Chamisso on a trip for
the study of sea fauna. In Berlin Nordmann published his
first work, MICROGRAPHICAL INFORMATION, dedicated to
the structure and taxonomy of parasitic worms (describing
many new forms, in particular "spainika" - Diplozoon paradoxwi) ,
and also the structure and development of parasitic copepods.16
In 1832 Nordmann was invited to join the department of zoology and botany of Lyceum Rishel in Odessa, and in the following
year he took the post of director of Odessa Botanic Garden
C128) . In 1833 Nordmann together with Rathke, S. S. Kutorgaya
and Steven travelled to the Crimea. In the following years
he travelled much in the south of Russia; in particular, he
led excursions in the Crimea of students of the Odessa
Lyceum. When he was sent with a scientific mission to Paris,
Nordmann visited with Milne-Edwards the coast of Normandy. In
1849 Nordmann came to Helsingfors University, where from
1852 to the end of his life he headed the department of
zoology. During the period of his scientific activity,
Nordmann published fifty-seven works in Russian, Latin,
German, French and Swedish on anatomy, embryology, taxonomy
and zoography of different groups of vertebrates (mammals,
birds, fish) and invertebrates (insects, spiders, crustaceans
and worms, mainly the parasitic molluscs, bryozoans and
Coelenteratae) ; he also studied botany and paleontology.
16. A. v. Nordmann, MIKROGRAPHISCHE BEITRAGE ZUR NATURGESCHICHTE DER WIRBELLOSEN THIERE (Berlin, 1832),
Vol. I, viii + 118 pp., 1: UBER BINNENWURMER IM AUGE
HOHERER THIERE; 2: BESCHREIBUNG EINIGER NEUEN
HELMINTHE. vol. II, xviii + 150 pp., 1: BESCHREIBUNG
EINIGER NEUEN PARASITICHEN ENTOMOSTRACEEN;
2: ERSTER BEITRAG ZUR NATURGESCHICHTE DER
LERNAEN.
In the previously mentioned MICROGRAPHICAL INFORMATION, the results of Nordmann ! s observations on the
development of parasitic copepoda Aahtheres peroanon appear,
and the larval stages of other related forms (Ergasilus
Sieboldi Nordm. , Traoheliastes polycolpus Nordm. and
Lernaeoaera cyprinacea) are described. These investigations
met the need to explain the systematic situation of parasites,
which at that time were related either to molluscs or to
the annulated worms, or even to coelenterates. If the relationship of some representatives of the mentioned groups Cf° r
example, Caligus) to crustaceans was to some extent only
probable, then these forms, such as Lemaeocera , which in
the adult condition are completely unlike arthropods,
remained in their systematic relations mysterious.
The embryonic development of Aahtheres was described
by Nordmann rather incompletely. "On the upper surface of
the yolk," he wrote, "is found at first a more transparent
region, an4 a granular part of the yolk, having the significance of a rudiment (KEIM) , turned into round or spherical
forms from which the more peripheral give material for the
formation of the rudiment membrane (KEIMHAUT) ... The
latter completely envelops the yolk and... forms later on
the wall of the body of the embryo (p. 78). After referring
to the separation of the head and the appearance of the rudiments of the extremities, Nordmann turned to the
characteristics of the nauplius larva, whose structure
becomes complicated after moulting.
Its central point was the description of the larval
stage of Lemaeooera oyprinaoea . The adult animals of
this genus were already known to Linnaeus; they are
characterized by a sacculated unopened body, deprived of
extremities and organs of sensations (Figure 37, A). "If
naturalists," Nordmann wrote, "are astonished at the
structure of the body of the mature animal of this kind,
their astonishment will be more natural when an opportunity
arises for them to observe the young animals. It can hardly
be imagined that there is anything more striking than an
offspring having absolutely nothing in common with its
parent. Before my eyes the egg receptacle in the mature
animal was ruptured, and the embryos set free. I saw young
animals exactly the same as I represent them in the drawing
(Figure 37, B) ; they have extremities, antennae, and even
bright red eyes" (128).
Nordmann 1 s observations met, according to him, decided
distrust from Berlin zoologists to whom he demonstrated
the nauplius larvae of Lemaeooera oyprinaoea. Later the
significance of this discovery was universally recognized
(129) .
After some years Nordmann published a small embryological
work concerning the Black Sea bryozoan Tenchca zosterioola A 7
In one zooid Nordmann found from four to seven eggs and
saw the penetration of the spermatozoids in the female cells
through an opening in their base. Later he saw the embryos
hatched from the eggs swimming by means of cilia and finally
settling in the seaweed Zostera. "As far as it is possible,"
Nordmann wrote, "I observed the transformation of the young
animals and the development of polyps from them." Greater
significance was possessed, however, by other embryological
work of Nordmann' s on the development of molluscs.
17. A. v. Nordmann, "Recherches microscopiques sur l'anatomie
et le developpement du Tendra zosterioola, espece de
polype de la section des Bryozoaires , " ANN. SC . NAT.
ZOOL., 11 (1838), pp. 185 - 191.
Figure 37. Lernaeooera cyprinacea (A) and its larva (B) (by Nordmann) .
The molluscs, especially the gastropods and lamellibranchs, are considered easily available material for the
study of development processes; therefore they early
attracted the attention of embryologists . The division of
the ova of the gastropod molluscs is described in the
investigations of Van Beneden and Windischmann, and the
division of the ova of the lamellibranchs — in the work of
Loven. In the ova of Modiolaria marmorata {Mytilus aisoors)
Loven observed a maturation division and described it, as
a process of ejection of nucleolus (embryonic spot) . The
important result of his work was the establishment of the
fact that during the division of the ovum of Modiolaria, the
separation in the vegetative hemisphere of non-nucleated
lobes takes place which soon merges with one of the blastomeres. This phenomenon is repeated many times during the
time of the following divisions. Loven saw clearly the
process of division itself, i.e. the deviation of the ovum
into separate blastomeres, but he did not trace the changes
in the nuclei, the equal and mutual distribution of blastomeres .
The separation of polar bodies in the ovum of the grey
slug is sufficiently clearly described in the above-mentioned
works of Van Beneden and Windischmann, although the division
following the separation of the polar bodies was inaccurately
described by them; they spoke about the formation of elevations on the surface of the yolk, divided by fissures. As
a result of this the entire surface of the ovum becomes at
the end as if embossed (BOSSELE) and resembles a raspberry,
The Belgian authors did not connect the formation of "the
yolk cells" with the appearance of the cuts on the surface
of the yolk; according to their concept, the yolk cells
originate under the superficial layer of the yolk. The
drawings illustrating their work show round bulges on the
surface of the developing ovum, and not its division as a
whole into separate blastomeres. The internal processes in
the divided ovum and blastomeres are neither described nor
illustrated in the drawings. Erroneous ideas of the processes
of division in the gastropodan molluscs were retained in the
work of 0. Schmidt appearing ten years later. 18 "The description of the ovum and the processes of division
inside it up to the formation of the embryo," Schmidt
wrote, "was given so completely by van Beneden and
Windischmann that I cannot add anything to it" (p. 278).
18. O. Schmidt, "Uber Entwickelung von Limax agvestis , " ARCH ANAT., PHYSIOL. (1851), pp. 278 - 290 .
The subject of Nordmann's above mentioned work on the
development of the gastropodan molluscs was the nudibranchiate
mollusc Tergipes Edwardsii . The description of the phenomena
of its embryonic development constitutes part of the extensive
monograph, 19 beginning with the presentation of anatomical
data. Nordmann described in detail the development of the
ovum in the ovary (5 28 - 31), characterized the yolk
elements of the ovum (§ 32) , the structure of the egg
ready for oviposition (5 33) and its membranes (5 34) . The
process of division {§ 39) of the ovum of Tergipes was
described by Nordmann as the following: the first fissure
of division can take place in different directions and
divide the ovum either into equal, or into very unequal
parts. The fissure dividing the ovum into four globules
of division takes place at right angles to the first
fissure, after which the ovum is divided into eight globules,
and so on. The generally uneven character of division is
noted in the following expression. "Although the tendency
to even progressive division cannot be disputed, it also
shows that the yolk, especially up to its acquisition of
the mulberry form, appears to divide very unevenly" (p. 573)
(Figure 38, a-g) . Nordmann noted especially that the fissures
cut all the mass of the yolk, i.e. that the division was
complete.
19. Nordmann, "Versuch einer Natur- und Entwickelungsgeschichte
des Tergipes E 'dwards ii , " MEM . , pres . a l'Acad. Sc.
St.-Petersb. par divers savants, 4 (1845) , pp. 495 - 602.
A brief translation of this work in French was published
by K. Fogt ("Essai d'une monographie du Tergipes Edwardsii,"
ANN. SC. NAT., 3, ser. Zool., 5 (1846), pp. 109 - 160).
All the course of development of Tergipes Nordmann summarized in the following statements:
1 . The chorion extend to one-fifth the diameter of
the ovum and becomes oval . 2 . At this time a light
fluid, similar to albumin, discharges from the ovum.
3. The yolk loses its spherical form, its mass
loosens, and the contours become wrinkled. 4. The
embryonic vesicle and the embryonic spot disappear.
5. The upper layers of the yolk lose their reddish
coloration. 6. In too many cases separate yolk
cells are separated from the other mass of the
yolk and give origin to the parasitic creatures. 20
7 . The ovum is divided by a fissure into two globules .
8. The division process continues regularly.
9. The yolk acquires the form of a mulberry.
10. A bubble of air leaves the yolk (?) . 11. The
surface of the yolk becomes granular. 12. First
establishment of the embryo. The yolk acquires an
elongated form, and then the form of a roughly
outlined triangle. 13. The distinct appearance of
the animal system and the cutaneous system. The
configuration of the embryo. 14. Twisting of the
anterior part of the embryo is noted (future
organs of movement) . 15. On the wide anterior area
folds appear, from which two lateral round growths
are gradually formed. The growths are transformed
into lobes and between them a third growth, the
foot, appears. 17. Beginning of the formation of
mantle and concha. 18. On the lobes the cilia grow.
19. The first movement of the embryo. On the foot
the vibrating cilia appear. 21. The lobes of the
sail become disc-shaped. The rotatory movement of
the embryo. 22. Cells sharing in the formation of
mantle are dissolved and disappear. 23. The concha
significantly enlarges. 24. The isolated cellular
rows indicate the formation of the attached muscles .
25. The formation of the internal organs, among
which the intestine is distinctly differentiated.
26 . The liver and other glandular bodies are clearly
seen. The anus and ganglia.
27. The cells forming the attaching muscles disappear. 28. Deposition of
the pigment in the eyes . 29 . Between the chorion and
the embryo the astonishing parasites appear to rush
out. 30. The complete formation of the embryo larva;
the roof of the shell opens and closes. 31. The
extension of the chorion. 32. The presence of the
young in the common ovum chamber. 33. The larva cuts
the chorion. 34. Hatching, (pp. 565 - 567)
20. Nordmann gave this parasite the name Cosmella hydrachnoides
and suggested the possibility of its origin from particles
of the yolk of Terg-ipes.
Figure 38. The development of Tergipes Edwardsii (by
Nordmann) a-g — division of the ovum; h — larva,
,36
From the above it can be seen that Nordmann's main
attention was to the relatively late stages of embryonic
development, while he investigated the process of division
of Tergipes only extremely incompletely.
The brief discussion of the works of van Beneden,
Loven and Nordmann on the embryology of molluscs demonstrated
the level of knowledge in this sphere at that time. When
the development of molluscs was studied by the Russian
embryologist N. A. Warnek, his work can be unconditionally
called classical (131). Warnek's scientific activity, unfortunately, was brief and was cut short by external circumstances,
Nikolai Aleksandrovich Warnek was born in 1821. When he
was eighteen years old he entered the faculty of law at
Petersburg University, but in the same year he transferred
to a department of the faculty of philosophy. During his years
of study, Warnek received in 1843 the gold medal for his
"Process of Moulting of External Tegmens and Formation of
Millstone in the Ordinary River Crayfish." In 1846 he graduated
from the university with a candidate's degree, and for three
years he taught botany and zoology in Gorn Institute. Warnek
received his master of science degree for his work on the
structure and function of the crayfish liver^l and in 1849
he started reading lectures to naturalists and medical men
in comparative anatomy and physiology in Moscow University,
first as junior scientific assistant and then as extraordinary
professor (from 1852) . In 1848 Warnek wrote the vast work
"On the Formation and the Development of the Embryo in the
Gastropodan Molluscs," which was published two years later
in the BULLETIN OF THE MOSCOW SOCIETY OF NATURALISTS; 22 a summary of this work was published abroad under
21. N. A. Warnek, "On the Liver of River Crayfish in
Anatomical and Physiological Relations" (SPb., 1847),
40 pp.
22. warnek, "Ober die Bildung und Entwickelung des Embryos
bei Gasteropoden," BULL. SOC . NATUR . MOSCOW, 23
(1850) , pp. 90 - 194.
the title "On the Process of Division and Structure of the
Ovum in Gastropods. "23
In 1860 Warnek retired and departed to Tver; where for
three years he was director of a secondary school, and then
director of the schools of Tver Province.
Until recently it was thought that Warnek 's scientific
and literary activity ended in 1863 and that in 1867 he died.
From recent work by T. P. Platovaya 2 ^ it is now known that in
1880 and 1881 Warnek read a report to the Moscow Society of
Naturalists on the biology of agricultural pests, and in
1884 a report on the microscopic structure of the fish ovum
and on the morphology and taxonomy of fish. The exact date
of his death remains unknown.
Concerning his retirement from Moscow University evidence
has been kept which is probably not completely objective and
in any case explains only incompletely the cause of his
retirement from teaching and from scientific work.
In the multi-volume apologetical biography of the
reactionary historian Pogodin, its author N. Barsukov25
reminisced about I. A. Mitropolsky, who was in 1850 a student
in the Faculty of Medicine in Moscow University. According
to Mitropolsky, when Warnek was reading zoology and comparative
anatomy he was noted for an extremely scornful nature; due to
this he was disliked by the students and medical men. One
of Warnek 's clashes with the students, Mitropolsky reported,
ended in a noisy scandal, in which both students and medical
men participated. In order to confirm to the reader the
ultimate rightness of the students, Mitropolsky declared that
"the students could endure the tricks of the professor only
if his lectures gave them anything useful. But they got
nothing from them."
23. Warnek, "Uber den Furchungsprozess und die Struktur des
Eies der Gasteropoden," FRORIEP ' S TAGESBERICHT ,
No. 280 (1851) , pp. 43 - 44.
24. See (131) .
25. N. Barsukov, ZHIZN'I TRUDY M. P. POGODINA (Life
and Works of M. P. Pogodin), Vol. 16 (1902), pp. 116 - 117
Another aspect of this incident is mentioned in the
reminiscences of L. V. Lebedinsky, which were published
in VOICE OF THE PAST in 1912.26 Lebedinsky characterized Warnek as talented, but very proud and sharp in his
treatment of people, who "excited against himself the
students and medical men by his tactless relations with
them and an unusual strictness during examinations."
Handbooks in Russian in zoology and comparative anatomy
were absent at that time, and Warnek suggested foreign
books, which the students, due to their ignorance of the
languages, found difficult to use. He did not read a
systematic course, but selected works of his own, which
he considered more important and interesting. In answer to
the protests of students on this occasion Warnek declared
that "the students of a university are not secondary schoolboys, they must work independently. The professor in his
lectures has only to point out a direction and a method by
which the students must carry out their work." The dissatisfaction of the students and medical men was expressed
in the organization of meetings in which the students of
other faculties shared, students of law and philologists
who did not attend the lectures of Warnek and could not
judge their effects and defects. In these meetings it was
decided to criticize the professor, and then oblige him
to leave the department. Only some of the students and
medical men, Lebedinsky continued, strongly defended the
professor. "They said that Warnek was regarded differently
and they loved him for his talented presentation of the
subject." The planned obstruction was nonetheless carried
out, mainly by students of other faculties who shared in
the action, especially those of the faculty of law. This
produced on Warnek a stunning effect. "Warnek went into an
adjacent room," Lebedinsky wrote, "where he fainted away; it is even said that blood gushed from his throat." Soon
after this incident Warnek retired, insisting on it despite
intensive persuasion.
26. UZ ZHIZNI MOSKOVSKOGO UNIVERSITETA. VARNEKOVSKAYA ISTORIYA. "GOLOS MINUVSHEGO" (From the life
of Moscow University. Warnek' s History. "Voice of the
Past") (Otd. ottisk, pp. 210 - 218). For this source, as
well as for the answer in HERZENOVSKY "K0L0K0LE" (see
[132]), the author expresses deep gratitude to V. V.
Sorokin .
Already this comparison of evidence shows the impossibility of placing complete responsibility on Warnek himself
for what occurred. The objective evidence of Lebedinsky about
Warnek f s talented lectures to the students and medical men
and their good relation to him unmasks the tendentious
assertion of Mitropolsky that Warnek' s lectures gave nothing
to the students. The impression one has of Warnek is of a
serious, exacting professor who loved his subject and taught
it on a high level. Certain students and medical men understood his efforts and valued his services to them, while
the majority of students were interested in the applied
questions and not in understanding Warnek' s theoretical
views, and saw him only as a strict examiner who shamed them,
in addition to his ridiculing of their ignorance.
In all this sad "history of Warnek" there is still one
significant side which can be read between the following
lines in Lebedinsky 's memoirs: "Even among the professors
there were people who sympathized with the students, the
professor of theology Sergievsky among them. This handsome,
somewhat fanciful, young orator was sometimes present in the
department. He unnoticeably and cleverly approached the evils
of the day, said a few words hinting at an excellent understanding of the students, and was zealously rewarded with
applause. In the time of the aforementioned event, one of
his lectures, which was full of these hints, ended with the
following significant words: 'Yes, this darkness does not
triumph over the world!"'
It is logical to ask why Warnek was not pleasing to
the professor of theology. What world of darkness and damage
spreading from Warnek 's department was frightening to
Sergievsky? To answer this question is not difficult. Warnek
was a convinced materialist, as is easy to ascertain from
a study of his basic work. It is improbable that in his
lectures Warnek did not touch upon questions of ideology,
which, of course, was used against him by Sergievsky, who
considered it his responsibility to protect the world of
"religion" from the "darkness" of the natural-scientific materialism. Sergievsky's Jesuitical activity made the best
use of the dissatisfaction of the protesting group of
students, who played a role obviously of considerable
importance in the organization of the obstruction, which
took place with the obvious support of the administration
of the university, usually so vigilant when the matter is
concerned with student disturbances (132) .
The fault, or rather misfortune, of Warnek was his
sharp and derisive character, his inability to adapt
himself to his surroundings. Warnek did not want to
reconcile himself with the manifestations of ignorance,
even if its carriers were respectable scientists. This
can be witnessed by Warnek 's review of a scholarly book
by A. Bogdanov. 27 This review shows great and extensive
knowledge and unquestionable educational talent of the
reviewer and his understanding of the problem of the teaching
of natural sciences. In addition, the review is written in
an overly particular and caustic tone; Warnek did not let
pass any mistake, any slip of the tongue, or any lame expression from the author of the book. It is easy to imagine what
reaction Warnek' s article could produce in Bogdanov. If
Warnek behaved similarly in the professional milieu of
Moscow University, and used to deride the lectures of his
colleagues, then he undoubtedly provoked against himself
not only a certain group of students, but also some professors.
Valuable evidence of Warnek 's high character as a lecturer
and scientific worker is contained in the words of I. M.
Sechenov : 28
Junior Scientific Assistant Warnek taught us zoology.
He read simply and clearly, dwelling mainly on general
signs applied in zoology departments, and the description of protozoa was prefaced by a long treatise on
the cell in general. This last study was built, however,
on unprepared ground. Moscow still did not think much
at that time of the microscope; among Warnek' s students
it was not used successfully, and in mockery they
nicknamed him "Cellular."
27. Warnek, "Zoologiya i zoologicheskaya khrestomatiya v obieme
srednykh uchebnykh zavedenii , " Author Anatolii Bogdanov,
OTD. PERVY ZH. MNP, ch. 118 (1863), pp. 47 - 73.
28. I. M. Sechenov, AVTOBIOGRAFICHESKIE ZAPISKI (Izd. AN
SSSR, 1945) , 176 pp.
In a footnote Sechenov added: "Too late, I have learned
that Warnek and the famous botanist Tsenkovsky were among
the first Russian biologists who worked at that time with
the microscope."
On Warnek' s socio-political opinions there is no
information. His statement against Eogdanov's book, based
on the principles of Darwin's evolutionary study, was
published in a government journal; objectively, it could
have played a reactionary role. The mutual relations of
Warnek with the students creates the suspicion of the
possibility that the rising conflict was of a political
nature. Concerning Warnek 's "political orientation," it
can be seen that he was later assigned responsible posts
in the Department of Education. A search of the archival
material may throw light on this subject.
For a characterization of Warnek' s scientific work and
world view, it is necessary to gain acquaintance with the
contents of his basic work, "On the Formation and Development
of the F:mbryo in the Gastropodan Molluscs. "^ As an epigraph
to this work Warnek selected the words of Reii: "The phenomena
of individual life are the necessary result of formation and
merging." This idea, as can be seen from the final chapter
of this work, Warnek interpreted materialistically. Warnek
himself also considered that only in the definite mixing of
substances in definite spatial position [form) can the
solution of vital phenomena be sought. Science, in his
opinion, is not in need of additional idealistic assumptions.
As it is seen from the introduction of his work, Warnek
considered his principal task to be the explanation of how
the yolk, i.e. ^substances of the ovum, are transformed into
tissue of the embryo and what conditions this transformation.
The solution to this question, in Warnek' s opinion, is possible
only through a thorough investigation of the processes of
development, and that is why he also outlined the following
vast program. First of all, Warnek suggested, it is necessary
to study the reproduction of molluscs, either hermaphrodites
or separate sexes. For this aim the following must be studied:
1) structure of male and female sexual organs; 2) origin of
29. See footnote 22 of this chapter
embryo, i.e. development of yolk (ovum) and semen;
3) processes occurring during copulation, i.e. the influence
of sperm upon the yolk, and finally 4) formation of the
additional parts of the ovum — albumin, membranes and ovum
cocoons .
Only after this it is possible to begin the study of
embryonic development, which Warnek divided into two periods.
The first period includes the development of the fertilized
ovum, i.e. the process of division and preparation of the
development of organs, and the second, the period of development of all systems of organs of the developing animal.
In his investigations Warnek proceeded from the following
situations established by the embryologists: 1) for the
transformation of yolk into an embryo fertilization is
absolutely necessary; 2) this last consists of the material
influence of semen on the yolk; 3) this influence takes place
only in an infinitely small space, therefore the spermatozoa
of the semen must come in direct contact with the mass of
yolk; 4) the result of fertilization is the formation of the
elementary organs of the embryo (cells); 5] the cells acquire
different forms, grouped in complicated organs of the embryo,
and thus form its body.
Warnek expressed regret that embryologists could not
completely solve the following important questions related
to the development of animals: 1) how the yolk, i.e. substance
of the ovum is transformed into tissue of the embryo; 2) how
its transformation at the time of development occurs; 3) in
what does the secret influence of the semen on the yolk
consist.
The first question Warnek considered to be solvable,
and he expressed the hope that in a short while the other
two questions would also be explained.
The solution of the principal questions of embryology,
in Warnek' s opinion, could take place on the basis of the
following presentations:
The cause of the phenomena conditioning the beginning
and the subsequent development of the embryo is
ordinarily attributed to the vital power, which absolutely clearly shows that the beginning of life
must be sought in the formation of the ovum in the
ovary, and the beginning of its development in
fertilization, namely in the influence of semen on
the yolk. And, of course, the essence of the powers
which condition all phenomena of nature remains
unknown. If all this can reduce the different
phenomena to one cause, this will make a great step
toward the goal we have established, (pp. 94 - 95)
During the last four years, aspiration to a solution
of the aforementioned general questions was my
dearest hope. Devoting myself year after year to
the study of the formation, development, and
functions of cells, I have stuck to the idea that
elucidation of the ideas of development and activity
of the cells is the only way to select from the
labyrinth of recent presentations about the organic
world. The explanation of the causes of the vital
activity in the cells leads to a clear presentation
about life in general and about causes directing it
throughout the organic world, (p. 95)
The following work of Warnek represents the first part
of investigations of his planned program; it is concerned
with ovum structure and processes of its division in the
gastropodan molluscs.
Warnek began with the description of the form of oviposition of the different fresh-water snails; he detailed the
periods of oviposition in the region of Petersburg and the
structure of gelatinous mass surrounding the ovum. Later he
described the structure of the laid ova, especially in the
species used in this work: Lymnaeus stagnalis and the slug
Limax agrestis . Transferring to the principal part of the
work, throwing light on the study of division, Warnek paused
at the characteristics of the yolk granules filling the
fertilized ovum. Among these granules Warnek saw a light
spot which was not delimited from the yolk membrane and was
always situated in the center of the ovum. All the process
of division of the ova of the molluscs which he studied Warnek
divided into stages, described in succession.
FIRST STAGE: For this stage, characteristic phenomena
are taking place in the above mentioned light spot, which,
in Warnek's words, occupies the place of the rudimentary
(embryonic) vesicle. This spot is in the beginning completely
round (Figure 39, 3), then becomes elongated and subsequently
takes the form of a biscuit and the shape of a figure eight
(Figure 39, 4 and 5). After that when the spot (i.e. nucleus)
is twisted by the means just described, it comes nearer to a
certain region of the periphery of the yolk (ovum) . The end
of the spot which is turned to the surface of the yolk widens,
and it acquires the form of a blunt rounded cone, in addition
between the outer area of this cone and the membrane of the
yolk a transparent crescent-shaped region appears (Figure 39,
7) . From this crescent-shaped region two small vesicles become
separated, which, being isolated from the ovum, remain near it
throughout the following development. The place of deviation
of these vesicles becomes the center of formation of fissures,
later dividing the yolk into two, then into four parts.
The separation of the vesicles Warnek described as
follows: on the external surface of the crescent-shaped region
a small elevation appears under the yolk membrane. It
gradually enlarges, acquiring the form of a spherical segment,
a hemisphere, then a complete globule, which is set on a
sufficiently thick stalk. Then this stalk becomes unlaced,
and the globule becomes free (Figure 39, 9). After the
formation of one vesicle the second one appears exactly as
the first (Figure 39, 9 and 10). Thus, in Warnek's observations, the crescent-shaped region separates the forming
vesicles from the light spot (i.e. from both nuclei of the
ovum) . He concluded that the nucleus does not share in the
formation, at least, of the external vesicles. He made this
erroneous conclusion because the intravital observations which
Warnek used did not suggest tracing the processes taking
place in the nucleus. From this came the further erroneous
claim that the separating vesicles could not be regarded as
the vesicles of Purkinje or its remnants. In accordance with
this assertion Warnek refused to recognize for the vesicles
separated from the ovum that important role claimed for them
by many authors, and he objected to the name "directing
vesicle" {vesioula directrix') . Warnek did not like this
name as it returned embryologists to the time when they
believed in an Archean spirit directing vital phenomena.
The presence of vesicles where the formation of fissures
of division begins did not prove, in Warnek's opinion, that
the topographical position of the fissures was determined
by the vesicles. Preferably, as he thought, the matter was
the contrary: the vesicles are separated where the center
of division is present. In later stages of division many
fissures appear without preliminary separation of the vesicles
After the separation of both vesicles the transparent
crescent-shaped region also disappears. In the ova of the
slug there appear two nuclei distinctly separated from each
other. At this time the nuclei each acquire distinct contour
and a large nucleolus.
THE SECOND STAGE begins with the loss of the membranes
of the nuclei and their merging into one common mass. This
mass acquires an extended form, situated on the longitudinal
diameter of the yolk, i.e. at right angle to the position
which is characteristic for the first stage (Figure 40, 11) .
Then the nucleus becomes biscuit-shaped (Figure 40, 12) and
at the same time the division of the yolk begins. In the
last description Warnek used topographical terms, and their
significance is explained in the literal remark. The transverse
diameter he called the diameter; passing through the vesicles
from the yolk at right angle to it the longitudinal diameter
is situated. The terminal points of these diameters he called
poles: the dorsal pole in the place where the external vesicles
were present; the ventral pole situated against the dorsal one;
the poles of the longitudinal diameter designated as right and
left.
The division of the yolk is preceded by a thickening of
the dorsal pole, in its field; then a fissure in the form of
a cut appears. Due to its deepening the ovum becomes in form
more like the kidney (Figure 40, 13) . The direction of the
fissure does not coincide with the transverse diameter; it
is inclined to it at a 45-degree angle. The light spot
decreases and becomes less noticeable even in the ova of the
slug, and in (Lymnaeus stagnali-s) it is not seen in the
majority of cases from the very beginning of the division.
Figure 39 . Warnek ' s drawings for his work on the development of
gastropodan molluscs (development of Lymnaeus
stagnalis)
3-5 — the division of the light spot in the fertilized
ovum;
7 — at the external area, light spot coming near the
surface; a light crescent-shaped formation appears; the
beginning of the formation of elevation;
the elevation acquires the form of a club-head;
9 the second elevation begins to form;
10 — the second vesicle separates; the light spot becomes
spherical; the crescent-shaped region disappears;
19 — completely separated yolk globules;
22— the beginning of secondary nearness of the yolk
globules ; between them a light space ;
25— the greatest nearness of the yolk globules; the
nuclei are not seen;
25a— no n- simultaneous division of nuclei into two primary
yolk globules;
27— non-simultaneous division of yolk globules;
28-30 — gradual distortion of first fissure of division;
3 4 cross -shaped position of the globules of division;
36a — no n- simultaneous division of the nucleus into four
yolk globules;
40— globules of division of the fourth stage (la- Id) are
situated in the space between globules of the third stage
CIA- ID) ;
43— extension of nuclei in the yolk globules of the
third stage (1A-1D) ;
45— the yolk globules of the fifth stage (2a-2d)
separated from the globules of the third stage (2A-2D)
and situated between the latter;
46a — adjacent situation of nuclei, showing the origin
of the yolk globules of the fifth stage from the globules of
the third stage;
47 — the yolk globules of the sixth stage (la 2 -Id 2 )
separated from globules of the fourth stage (la 2 -Id 2 );
50 — the ninth stage of division.
Figure 40. War-nek* s drawings for his work on the development
of gastropodan molluscs (development of a slug)
11 — in the place of the lost membrane of the nucleus,
the oval light spot is seen;
12— the yolk has the form of a globule thickened from
one side; the light spot is elongated;
13 — in the thickened side of the yolk the beginning of
the formation of a fissure is seen; the light spot becomes
more elongated;
14— the fissure twists the yolk diagonally; the light
spot is extended;
15 — the fissure envelops half of the periphery of the
yolk; at the ventral pole is the beginning of the formation
of a fissure in the form of a deepening;
16 — deeper twisting of the yolk, viewed from the dorsal
pole;
17 — the yolk globules are completely separated from each
other; the light spot is hardly noticeable;
26 — the nuclei are deprived of membranes and begin to
extend in a direction perpendicular to the longitudinal pole;
26a — still more distinct changes, the beginning of which
is illustrated in Drawing 26;
29a— the stage of four yolk globules.
On the basis of experiments on ova which had been
pressed in water, Warnek concluded that the structural
changes in the nuclei depend upon the change of their
chemical composition. The chemical changes that appear,
in Warnek' s opinion, condition also the further transformations in the developing ova. After that when the fissure
begins in the dorsal side (Figure 40, 14), it passes around
half the periphery of the yolk; also in the ventral side,
a deepening appears (Figure 40, 15) and the yolk is twisted
by the meeting fissures. It acquires first the form of a
biscuit, and then two united or even completely isolated
globules (Figure 40, 17) . At the time of division of the
yolk the light spot (nucleus) divides into two parts, each
of which at first have caudiform processes, directed to the
point of contact of the yolk globules (Figure 40, 16) . The
processes quickly disappear, and the spot becomes spherical.
These phenomena, according to Warnek, characterize
the first half of the second stage of division. At this
time it is not possible to isolate the nuclei by (pressing)
the yolk globules; from which it must be concluded that the
nuclei are still deprived of membranes,
The second half of the second stage begins with the
dividing globules moving nearer. Between them a noticeable
cavity, formed from a transparent substance, emerges . Warnek
considered this transparent substance the product of separation of the yolk globules. At the time the dividing globules
are moving closer, the contours of the nuclei in them again
become clear, i.e. the membrane appears (it is clearly
noticeable in Li-max) by strongly refracting the color of the
nucleolus. In Lymnaeus stagnalis the nuclei, at first, are
situated near each other (Figure 39, 22), then separate,
sink in the depth of the yolk globules, and at the end
become invisible (Figure 39, 25) . Following the described
phenomena each of the two yolk globules become pear-shaped.
This outlines the passage to the third stage of division,
during which the yolk is divided into four parts.
THE THIRD STAGE begins with changes in the nuclei,
acquiring a biscuit or figure-eight shape (Figure 39, 25a);
in the drawing it is seen that their division is not
accomplished simultaneously: when one divides, the other keeps the form of a quadrant. The details of these changes
can be traced only in the transparent ova of slugs. When the
yolk takes the biscuit form, the nuclei are elongated
(Figure 40, 26); following this the elongation and twisting
of the yolk globules themselves takes place (Figure 40, 26a) .
The twisting in Lymnaeus stagnalis and slugs begins in one
globule earlier than in the other (Figure 39, 27), however,
soon after this difference smoothes out. The boundary between
the globules of division, corresponding to the first fissure,
is at first straight (Figure 39, 27), and then becomes
curved (Figure 39, 29, 30), and the yolk globules are situated
crosswise in two planes.
The processes taking place in the nuclei were described
by Warnek as follows: "The membranes of the nuclei disappear,
the nuclei elongate, take an oval biscuit-shaped form, then
bulge out, and finally each nucleus from the beginning of
the division of the yolk globules is divided into three parts.
From these parts of the nuclei only four are present in the
globules of the division, and two gradually disappear in
the fissures between the globules. The four nuclei at first
have the form of a comet. When the division of the yolk
globules is finished, the CAUDIFORM processes of the nuclei
extend and the nuclei again acquire the rounded form"
(pp. 146 - 147). The three parts into which, according to
Warnek, each nucleus is divided correspond to the two daughter
nuclei and to the achromatic figure of mitosis situated
between them.
The final step of the third stage is the formation of
the membranes around the nuclei and subsequent turning of
the cross-shaped globules of division. Two of them are in
contact with each other on the dorsal side, and the other
two on the ventral (Figure 39, 34) . Between the yolk globules
a rhomboid space appears during which this is especially clear
in Limax.
THE FOURTH STAGE . At the beginning of the fourth
stage the nuclei in the yolk globules of Lymnaeus stagnalis
again become unnoticeable from outside. During the crushing
out of ova it is possible, however, to see the changes
occurring in the nuclei, which, as in the previous stage
do not take place at the same time. In Figure 39, 36a it is seen that in the two globules of division which have the
longitudinal form, one nucleus is biscuit-shaped, and the
other consists of two isolated parts. After that, when the
nuclei become elongated (in slugs this is seen also in the
intact globules of division] , each yolk globule stretches
and becomes pear-shaped. Then the twisting occurs in this
form so that the newly forming globules are of unequal size,
each separating a region of one third the size. The four
smaller globules become displaced and are situated in the
spaces between the larger two (Figure 39, 40) . "The remarkable
reciprocal situation of the yolk globules," Warnek wrote,
"is kept and is repeated in all the following stages; this
allows one, without ever noticing the further formation of
the yolk globules, to solve the question, what globules of
division result from each present globule. During this it
is necessary to keep in mind the position of the nuclei and
the relative size of the yolk globules" (p. 153).
These accurate observations surpassed those investigations of nearly a quarter of a century afterwards. By the
initiative of A. 0. Kovalevsky, the blastomeres of the
dividing ova were given individual designations (in letters
and numbers) , tracing during the process of development the
fate of each blastomere and its derivatives. Warnek formally
applied a less suitable and obvious method of designation
of the globules of division and their descendants. He
named the blastomeres arising in one or the other stage by
the number of this stage, keeping for them the same designation also in the following stages of division. During this he
mentioned that the yolk globules changed from stage to stage,
so that, for example, during the transfer to the fifth stage
the globules of the fourth stage were already unequal to the
globules of the fourth stage at the moment of their formation.
They decrease in size and are changed by chemical properties
and internal structure.
THE FIFTH STAGE. In this stage the formation of
the new globules of division follows the rules which also
hold for the following stages of division. Instead of the
sixteen yolk globules which must be present if each of the
eight globules of the fourth stage is divided, here only
twelve globules are found. This can be explained by the fact
that during the fourth stage only four large globules are divided, and the other four small globules remain unchanged.
Before the division itself the nucleus of the ventral (large)
yolk globules become invisible, although on crushing out of
the ova, it is seen that they elongate, i.e. they are present
in a condition of division, while the nuclei or the dorsal
(small) globules are not divided and remain round (Figure 39,
43) . The dividing yolk globules are stretched and twisted
in the diagonal direction (Figure 39, 45). The newly arising
yolk globules (again smaller in size than those which gave
them the origin) are generally situated by the general rule,
in the spaces between the large vegetative globules. The
twelve globules of division present in the fifth stage are
situated in three rows. The ventral row consists of four
globules of the third stage, in the dorsal side four yolk
globules of the fourth stage are present, and between the
ventral and dorsal globules four newly arising yolk globules
of the fifth stage are situated. The globules of the upper
and lower rows stand against each other, and in the spaces
between the globules of these rows the globules of the middle
row are present. Concerning the origin of globules of the
fifth stage from the globules of the third stage it is
judged by the neighboring situation of their nuclei (Figure 39,
46a). In the second half of the fifth stage, as in the previous
one, the smoothing of the surface of all globules of division
and their nuclei becomes more distinct. Between the yolk
globules a vesicular light space appears.
THE SIXTH STAGE. At the beginning of this stage four
yolk globules of the fourth stage become more convex and
the globules of the sixth stage separate from them. The
yolk globules of the fifth stage remain the spaces between
the globules of the third stage (Figure 39, 47). The total
number of globules of division in this stage is sixteen.
THE SEVENTH STAGE is characterized by the three
divisions of the globules of the third stage, giving rise to
the four yolk globules of the seventh stage. The total number
of globules of division is twenty.
THE EIGHTH STAGE. In this stage the four globules of
the eighth stage are separated from the yolk globules of the fourth stage. The total number of yolk globules grows to
twenty- four.
SUBSEQUENT DIVISION. In each of the following
stages four yolk globules are formed. As an example the
ninth stage can be employed. In the given drawing (Figure 39,
50) the dividing ovum is pressed, so it is possible to see
a great number of globules of division. In the middle the
yolk globules of the fourth stage are situated. To these
last the oldest and large globules of the fourth stage are
adjoined, moving far aside from each other by pressure.
To the left of the globules of the third stage the globules
of the fifth stage are present, obliquely from which the
youngest globules of the ninth stage are twisted; they are
situated in the spaces between the globules of the fifth
and seventh stages. The globules of the seventh stage
originate also from the globules of the third stage and
are situated to the right of the last globules. Finally
between the yolk globules of the fifth, seventh and fourth
stages the globules of the eighth stage are present, arising
as a result of a second division of the globules of the
fourth stage.
Further Warnek carried out an analogical analysis of
the fifteenth stage and established the origin of all globules
present at this moment of division.
It is instructive to compare the genealogy of blastomeres,
established by Warnek, in gastropodan molluscs with recent
data. This comparison shows the complete agreement of Warnek 's
results with recent data, as seen from the table. In it the
designations applied by Warnek and the presently applied
literal numerical designations are given.
The same comparison is given in Figure 41, where the
contours of drawings 40, 45, 47 and 50 of Warnek are repeated
and the data of the recent designations of blastomeres and
their designations by Warnek are compared.
There is no doubt that Warnek completely and distinctly
chose to follow the fate of the separated blastomeres ("yolk
globules") and the participation of their descendants in the
formation of organs of the developing animal. Selecting an irreproachable method by which this problem can be solved,
Warnek in the first published investigations on the
embryology of gastropodan molluscs described the first
period of development up to the formation of the spherical
multicellular stage, i.e. the blastula. Only at the end did
he briefly mention the following period, when "some yolk
globules share in the formation of first internal organ —
the yolk sac." This first internal organ, the yolk sac, is
of course nothing other than the endoderm of the embryo.
Concerning some details of division, Warnek noted that
during the division of the nuclei of the yolk globules the
nucleolus appears earlier than the nucleus when divided into
two parts, therefore it is possible to find nuclei with two
nucleoli. Warnek never saw the process of division of the
nucleolus itself. The division of the nucleus in his experience
was always accomplished by one plan, which in the early
stages of divisions was the same as in the subsequent development of the embryo. This division in all conditions takes
place after the stage of stretching of the nucleus, which
then acquires the shape of a biscuit and a figure eight and
is finally transformed into two separate nuclei.
The globules of division Warnek identified as the
elementary organs, i.e. cells, and considered that their
multiplication, beginning at the time of division, continued
throughout the period of development and even through the
entire life of the animal.
All the activity of the developing embryo and the
animal forming from it is, in Warnek' s opinion, the result
of that primary influence which the ovum ["yolk mass")
is subjected to by the semen. "This influence," Warnek
wrote, "has a purely chemical nature; therefore the explanation for this is still obscure for us; the vital phenomena must
be given by physicists and chemists" (p . 168). "The effective
element in the organism," Warnek continued,
is the material; this same material influences also
outside the organism. If we explain this activity by
chemical and physical powers, then there is no reason
to deny the activity of these powers in the organism
as well. Although these powers still cannot be completely explained, we do not possess the right
to discard them and resort to the help of this power,
which exists only in the imagination. Can we explain
the phenomenon of crystallization? Why does sodium
chloride always crystallize in the form of a cube,
and pure carbon in the form of an octahedron? Is
not the formation of globules of division, from the
point of view of form, a kind of crystallization of
organic matter? The successes of organic chemistry
belong to us, because the processes accomplished
during nutrition, respiration, and excretion are
more satisfactorily explained by means of physics and
chemistry than by means of a special vital power. This
power has retreated into the dark field of our
knowledge about the functions of brain and nerves and
still dominates in the sphere of embryology. However,
new histological directions make the study of the
vital power even more unsteady in this sphere, so we
are not far from the time when chemistry will
completely exclude it from there as well. The concept
of vital power must remain as a reminder of our
previous ignorance. Only quite recently the influence
of semen on the yolk was called dynamical; this
expression shows only that the phenomena of fertilization could not be explained, (p. 170)
In these words Warnek exhibited the materialistic world
view with complete clarity. He decisively objected against
the dull idea of the vital power for the explanation of
phenomena of organic life. The only way in which this
explanation could be achieved Warnek considered to be the
physico-chemical investigations of vital phenomena. Regarding
fertilization as a chemical process, Warnek thought that
the subsequent transformations of the dividing ovum have as
their source continuous chemical changes. Of course, Warnek' s
materialism has a mechanical character, but it is not
excessively simplified vulgar materialism.
Warnek 's embryological opinions are expressed in his
theses, the most important being:
The yolk mass after fertilization undergoes chemical
changes, therefore the fertilization itself must be regarded
as a chemical process. It causes changes in the unfertilized
ovum which are necessary for further development of the embryo
Table
The Genealogy of Blastomeres
Comparison of Warnek's designations— stage numbers (in parentheses! in comparison with the recent literal-numerical. Cln
the square brackets the designation of the resulting blastomeres of the previous stage are repeated.)
Key:
1 . Stage of division by Warnek
5. The sixth
2. The third
6 . The seventh
3. The fourth
7. The eighth.
4. The fifth
Figure 41. Comparison of Warnek's designations of blastcraeres
(stage numbers) with the recent literal-numerical
designations of blastomeres . These and the other
designations are put in the contours of Warnek's
drawings .
During the development of the embryo, further changes of
chemical processes take place.
The gastropodan molluscs are characterized by complete
division.
The globules of division may be considered true cells.
In each stage of a division process four yolk globules
are formed, i.e. the division proceeds not in geometrical,
but in arithmetical progression.
Beginning with the third stage, the globules of division
have unlike sizes.
Warnek's work produced a new page in embryology,
directing the investigations of the history of individual
development towards the study of subsequent changes of the
fertilized ovum and the forming from it of blastomeres, and
towards a study of the fate of the separate blastomeres and
their descendants during the subsequent formation of the
embryo. In this sense Warnek's investigation foreshadowed
the works of A. 0. Kovalevsky and his countless followers
who were studying either descriptively or experimentally the
transformation of the elementary organs of the dividing ovum
the blastomeres into systems of organs of the forming organism
N. A. Warnek was for a long time undeservedly forgotten.
His classical work is rarely cited and not always mentioned
even in the embryological summaries and textbooks, although
he unquestionably deserved a place of honor in the history
of Russian and world embryology.
The investigations of Grube, Nordmann and Warnek were
monographical descriptions of the embryological development
of certain representatives of the invertebrates. These works,
with all their significance, did not answer, however, the
requirements of comparison of the phenomena of development
in different types of animals. The first attempt to include a
wide number of invertebrates in embryological investigations
was done by A. Krohn, whose services in this sphere are much
undervalued. Krohn was so thoroughly forgotten that his
name was not mentioned either in the encyclopaedias or in
the biographical reference books. 30 The following circumstance
is sufficient to attract the attention of historians of Russian
science to Krohn.
30. For help given in researching biographical and bibliographical data about Krohn, the author thanks the
biological section of Saltykov-Sedrin Gos. Publichnaya
library in Leningrad, especially librarian V. L. Levin
During the first committee discussion of Baer's prize of
the Russian Academy of Science in 1867, considering possible candidates for the prize, the following was stated: "If the
matter concerned the crowning of previous scientific works,
then the committee does not doubt that the prize belongs to
one of our compatriots, Krohn, who was born in Petersburg.
For many years from the fertile shores of the southern seas
he collected a rich material which he investigated for the
development of different animal forms. His investigation
resulted in many excellent works which deserve respect from
the scientists of all countries. However, the competition
was to take under consideration only the works of the last
three years. "31
The absence of biographical information about Krohn is
compensated for by some bibliographical data. It is established that Krohn published no less than eighty works, 32
including some small monographs. Many of his publications
were accompanied by indications of the time and place of
performance of the corresponding work. With these indications
one can form opinions about the life of Krohn, who spent no
less than thirty years in travel with the aim of scientific
investigations, zoological and embryological .
31. "Extract from the report of the committee on the discussion
of the prize of secret adviser K. M. Baer, read in public
meeting of the Academy of Science on February 17, 1867 by
Academician Ovsyannikov, " NATURALIST (1867), Vol. 4,
No. 7-9, pp. 98 - 104; No. 10 - 12, pp. 146 - 148. The
extract cited is on p. 99.
32. The list of publications by A. Krohn is presented in the
following :
1. CATALOGUE OF SCIENTIFIC PAPERS, compiled by
the Royal Society of London (Vol. Ill, 1869- Vol. VIII,
1879) ;
2. w. Engelmann, BIBLIOTHECA HISTORICO-NATURALIS,
Suppl . Band: J. V. Carus und w. Engelmann, BIBLIOTHECA
Z00L0GICA: VERZEICHNIS DER SCHRIFTEN UBER
Z00L0GIE, WELC'HE IN DEN PERIODISCHEN WERKEN
ENTHALTEN VOR JAHR 1846 - 1860 SELBSTANDIG
ERSCHIENEN SIND, Ed. I - II (Leipzig, 1861), 2144 pp.
33. Information on the dates of Krohn' s birth and death are
taken from the brief bibliographical catalogue published by
the Library of Congress in Washington.
August David Krohn was born in Petersburg in 1803.33
Concerning his birth and student years of study of Krohn we could not discoyer any information. One of his early works
was produced in Vienna (1836); of the next work there is a
memorandum, Petersburg (1 83 7) , Later on Krohn was in
Heidelberg (18:59], and from 1840 he worked nearly continuously on the shores of the Mediterranean Sea and the islands
of the Atlantic Ocean. In 1840 and later he was in Naples;
from the autumn of 1844 to the spring of 1845 in Messina; 1848
in Nice; 1850 in Naples. In the beginning of 1835 and the
winters of 1853/54 and 1856/57 he again was in Messina; in
the winter 1855/56 and the spring and summer of 1865 he
worked in Funchal (in Madeira) and in Santa Cruz (Tenerif e) .
In December 1860, in May 1861 and in 1867 Krohn was in Nice,
and in the first half of the year 1869 he was in Naples. In
the intervals between travelling he lived in Paris (winter of
1851/52, spring of 1857) and in Bonn (summer months of 1851,
1853, 1855, 1857 and 1859, winters of 1859/60, 1864/65 and
1856/66, and also the second half of the year 1869) .
Concerning the last twenty years of Krohn 1 s long life of
eighty-eight years there is again no information.
During his travels Krohn maintained contact with his
country, as seen by the report of the conference of the
Academy of Science in Petersburg, which presents the
following records:
"Mr. August Krohn is a doctor who is famous for his works
on anatomy and physiology. He has sent the Academy a significant collection of invertebrate sea animals which was collected
by him near Naples, which, in quantity and quality, deserve
the thanks of the Academy" (Report from December 16, 1842) .
"Dr. Krohn sent again, as a gift to the Academy Museum,
two new collections, about one hundred species of fish,
crustaceans and others" (Report of September 13, 1844).
On the 7th of November 1855 Krohn was recommended as
candidate for corresponding member of the Academy of Science
in Petersburg, hit was not elected. The biographical data
given during this presentation shows only that he was born in
Petersburg, lived abroad, and wrote about thirty valuable works
dedicated to molluscs (IPaludina, Phyllirhoe and oephalpoda) ,
worms (SipunculuSj Syllisj Aloiopa) , and tunicates
{Doliolum) , 34
During his travels Krohn entered into friendly relations
with many great zoologists of the time, such as Johannes
Muller, M. Sars, and Delle-Kyaie, and also the young
investigators A. Kolliker, 35 k. Gegenbaur, 36 an( j a. Schneider.
Johannes Muller (1801 - 1851) was a great German zoologist,
embryologist and physiologist. For a long time he was the
editor of the widely distributed journal ARCHIV FUR
ANATOMIE, PHYSIOLOGIE UND WISSENSCHAFTLICHE MEDIZIN,
in which Krohn published about thirty articles. Part of this
information Krohn sent to the editor of the journal in the
form of letters, containing information about his last works,
and Muller published them in his ARCHIV which he sometimes
accompanied by remarks and additions, always with a friendly
and positive tone. In those remarks of Mill Her' s, discussions
can frequently be found revealing his high regard of Krohn' s
scientific activity.
Krohn willingly related his observations to the zoologists
who were working at the same time with him along the sea coast
C133) . His objective was to verify his data and to confirm
their authenticity, and equally to help also the beginning
investigators. His description of the planula hydromedusa
Cladonema, Krohn accompanied with the remark that he showed
them to Sars and Gegenbaur, thus certifying the accuracy of
his observations.
34. Archives AN SSSR, fund 2, inventory 17, No. 6. The author
is deeply grateful to B. E. Raikov. On a commission from
him, extracts of reports of conferences of the Academy
of Science were carried out here and an Archives Certificate
received.
35. "For accurate information about the structure of marginal
bodies in medusa," Krohn wrote, "I am grateful to my young
friend Kolliker from Zurich" (A. Krohn, "Einige Bemerkungen
und Beobachtungen uber die Geschlechtsverhaltnisse bei den
Sertularinen," ARCH. ANAT . , PHYSIOL. (1843), pp. 174 181} . Albert Kolliker was a well-known histologist and
embryologist .
36. Karl Gegenbaur (1826 - 1903) later became a famous
comparative anatomist and embryologist.
Anton Schneider (1831 - 1890) was a famous German
zoologist, who in a work on the development of the mollusc
Phyllirhoe bucephalumS? warmly mentioned "the repeated
friendly directions" which he received from Krohn not only
during the observations taken for that work, hit also
throughout the time of their presence together in Messina
in the spring of 1858. Significantly later, in 1867, Krohn
met with Schneider in Nice; Schneider was interested there
in larvae of any polychaetes which were covered with peculiar
porous membrane. Schneider found that these larvae were well
known to Krohn. In addition, as Schneider wrote, "Krohn,
with his characteristic generosity, gave me the relevant
pages of his journal that I might use the information
contained in it at my discretion, expressing the hope that
I could trace the further development of these larvae" (p. 498,
footnote) .
Other works reveal that Krohn did not intend to publish
his materials on the development of this polychaete, and
agreed that Schneider would do this himself. Schneider wrote
the work, its first part (description of the early stages)
containing his own materials, and the second the results of
Krohn 's observations on subsequent development. The work was
published under the names of both authors, Krohn 's name in
first place. 38
Helping to increase his material, Krohn showed at the same
time extreme punctiliousness in relation to the strange data.
This is illustrated by Krohn' s following remark on one of the
early works about the structure of the nervous system in the
echinoderms,39 as he sought to eliminating all shades of
suspicion of incorrectness in relation to the published data
of other investigators:
37. A. Schneider, "Uber die Entwickelung der PHYLLIRHOE
BUCEPHALUM," ARCH. ANAT . , PHYSIOL. (1859),
pp. 35 - 37.
38. A. Krohn und A. Schneider, "Uber Annelidlarven mit porosen
Hullen," ARCH. ANAT., PHYSIOL. (1867), pp. 498 - 508.
39. A. Krohn, "Uber die Anordnung des Nervensystems der
Echiniden und Holothurien im Allgemeinen," ARCH. ANAT . ,
PHYSIOL. (1841), pp. 1 - 13.
After I finished my observations on the nervous
system of echinoderras and reported all the existed
to Mr. Delle-Kyaie, I learned from this scientist
that Mr. Van Beneden, a year before, had already
discovered traces of the nervous system in echinus,
information about which had appeared in L'INSTITUT.
Because I could never get the proper issue of this
journal, I should not be birred for not mentioning
Van Beneden' s observations . (p. 7)
In all his works Krohn, with exceptional honesty and
modesty, mentioned the results of the work of his predecessors, not fearing to recognize the superiority of foreign
observations over his own. 40
No information was kept on personal events in Krohn 's
life. He was not connected in his work either with scientific
institutions or with universities. His life as a travellernaturalist hardly assisted the acquisition of a family of
his own. To his relatives Krohn superficially referred in a
letter to Johannes Muller:41 "After eight months absence,
during which I spent April and May in Santa Cruz in Tenerife,
I returned to Europe. The immediate cause for this was a
forthcoming meeting with close relatives, whom I had not
seen for some years" (p. 515).
In the first period of scientific activity (up to 1846)
Krohn' s scientific interests were concentrated in the anatomy
of vertebrates (fish, amphibia, birds) and invertebrates
(coelenterates, annelids, arachnids, chaetognatha, molluscs,
bryozoa, crustaceans, echinoderms, tunicates) . Incidentally
to his morphological investigation, Krohn found parasites
in the venous sinuses of cuttlefish (apparently diciemid) and
described new species of pteropod and cephalopod molluscs.
Zootomical and zoological investigations were continued by
Krohn, investigating the structure of protozoa, siphonophora,
40. Concerning the budding of the complex ascidian, Krohn
wrote: "Mechnikov, with greater success than I, has
traced the gradual development of buds" (Krohn, "Uber
die Fortpflanzungsverhaltnisse bei den Botrylliden, "
ARCH. NATURGESCH., 35 (1869), pp. 190 - 196).
41. ARCH. ANAT., PHYSIOL. (1856), pp. 515 - 522.
annelids, sea spiders, and arachnids, and describing new
species of annelids, chaetopods, and gastropodan molluscs.
There is much authoritative evidence on the accuracy of
Krohn's observations and morphological descriptions. These
are sufficient to justify the opinion of Kovalevsky, 42 who
was shared in the discussion of the nature of the so-called
"ventral saddle" of sagitta. Krohn in 184443 considered
this formation to be due to the nervous ganglion. Later,
W. Busch44 corrected Krohn's opinion, and, in spite of the
latter 1 s objection, Busch shared this point of view with
Keferstein, R. Leuckart, Pagenstecher, and K. Gegenbaur.
Keferstein did not agree with Krohn, but he gave credit to
his anatomical investigations: "Krohn, as it is known,
related this saddle's very great size to the nervous ganglion.
I, together with Busch, do not doubt in that this excellent
investigator was in the present question mistaken. "45
Kovalevsky, again investigating the anatomical structure
of sagitta, strongly supported Krohn against the above mentioned
authoritative zoologists. "I am against the new investigators,"
Kovalevsky wrote, "in considering Krohn correct concerning
the ventral ganglion, and I hope to convince my readers of
this also" (p. 135). "The ventral ganglion," he continued,
"has the form of a long oval or quadrangular body with four
large nervous trunks, from which two on the anterior end
continue to the brain or the cephalic ganglion, connecting,
as Krohn showed correctly, with the lateral nerves of the
cephalic ganglion" (p. 136) .
42. A. O. Kovalevsky, "Embryologicheskie issledovaniya chervei
i chlenistonogikh" (Embryological investigations of worms
and arthropods) (1871), IZBRANNYE RABOTY (Izd. AN SSSR,
1951) , pp. 123 - 266.
43. A. Krohn, ANATOMISCH-PHYSIOLOGISCHE BEOBACHTUNGEN
OBER DIE SAGITTA B I PUNCTATA (Hamburg, 1844), 16 pp.
This work was published a year later in French (ANN. SC .
NAT., 3 Ser., Zool., 3 (1845), pp. 102 - 116) and in
English (ANN. NAT. HIST. 16 (1845), pp. 289 - 304).
44. w. Busch, BEOBACHTUNGEN UBER AN ATOM IE UND
ENTWICKELUNG EINIGER WIRBELLOSEN SEETHIERE
(Berlin, 1851), viii + 143 pp.
45. Cited in the article by A. O. Kovalevsky.
From 1846, Krohn frequently turned to the study of the
phenomena of reproduction and development of different
invertebrates - coelenterates, worms, molluscs, crustaceans,
and mainly the echinoderms and tunicates.
The development of coelenterates is described in the
following words — first, concerning the hydromedusa Cladonema
and its\ development from the polyp Stauridium^ : the polyp
forms buds from which the medusae are formed. Similar to
oceanids in these medusae, as in Ooeanidae , in the walls of
the stomach the sexual products develop. If mature males and
females are situated in separate vessels, then after a short
time on the bottom and walls of the latter ova can be seen,
covered by a closely adjacent yolk membrane. That these ova
are fertilized, Krohn judged by the absence of any embryonic
vesicle and embryonic spot (nucleus and nucleolus) . Krohn
mentioned I later the process of ovum division, though not
describing it in detail, and referring to the fact that this
process was observed already by Dujardin, who did not, however,
evaluate/ its significance. Within two days after fertilization
the formed larva is seen in the egg, which later on leaves the
ovum membrane and swims with the help of cilia. The larva
was characterized by a light superficial layer and included
a dark, probably hollow nucleus (Figure 42, A). By its structure
the larvae of Cladonema are similar with the young of higher
organisms such as planulae (Aurelia, Cyanea, Cephea) . After
two to five days the planula Cladonema becomes rounded,
situated on the bottom, loses its cilia, and is transformed
into a disk, not changing its internal structure. In the
middle of the disk appears a round, hollow hillock, which grows
later into a cylindrical process, composed of two layers
present in planulae. On the upper end of the cylinder (rudiment
of the polyp) four hillocks form, corresponding to the
external ends of the future antennae. Already at this stage
the first stinging capsules are seen (Figure 42, B) . Thus,
Krohn concluded that "Stauridium resulted from the budding
of the medusa Cladonema, which reproduced by ova; the young
developing from the ovum is again transformed into the form
of a polyp. The subsequent change of heteromorphic generation,
from which more highly organized medusa develop must be
regarded as a generic form and is considered, consequently,
factually proved" (pp. 425 - 426).
46. A. Krohn, "Uber die Brut des Cladonema radium und deren
Entwickelung zum Stauridium," ARCH. ANAT., PHYSIOL.
1853, pp. 420 - 426.
A preliminary report on these observations Krohn included
in a letter addressed to Miiller.47
Figure 42. Planula hydromedusa Cladonema (A) and the
polyp developed from it Stauridium (B) by
Krohn .
Two years later Krohn published a report about the
structure of the early stages of development of the medusa
Pelagic. noctitucaA^ At first he found near Messina invertebrate
medusae similar to ephyra scyphomedusa, separating from "polypform helminths" (scyphistoma) , and he found also earlier larvae.
47. A. Krohn, "Uber einige niedere. Thiere. Brief liche
Mitteilung a. d. Herausgeber, " ARCH . ANAT . , PHYSIOL
C1853) , pp. 137 - 141.
48. A. Krohn, "Uber die friihesten Entwickelungsstufen der Pelagia nootiluca,"
pp. 491 - 497. ARCH. ANAT., PHYSIOL. (1855),
The supposition that they were stages of development of
Pelagia noctiluca was completely confirmed. After many
unsuccessful attempts, Krohn could carry out artificial
insemination. As a result of division ,larvae formed having a
cylindrical, usually stretched form (Tigure 43, A). The end
of the larvae (a) which is directed forward while swimming
is rounded, and the other (b) is chipped off. The surface of
the larva is covered by short cilia. In the blunt end occurs a
depression with an extremely small, round orifice is seen.
This orifice is the mouth, which leads to the round, clearly
outlined cavity of the stomach (c) , occupying the posterior
third of the body. Tne mouth and stomach, in Krohn 's words,
are already clearly differentiated already in the natural forms,
but it is still a non-hatched embryo. However in the present
stage the stomach is shorter and more rounded than in the
free larvae.
Mechnikov rated highly this discovery by Krohn. In
his monograph EMBRYOLOGICAL INVESTIGATIONS ON MEDUSA
(1866), 49 he noticed the weak interest in embryology by the
zoologists of the mid-nineteenth century. He wrote:
Even important generalizations, such as the similarity
between the two layers of coelenterates and the
embryonic layers of the higher animals, emphasized by
Huxley, and significant facts, such as Krohn' s discovery
of the formation of a stomach in pelagia by a
stretching of the blastomeres, remained without
attention and in a lower plane, (p. 284)
The correctness of these observations by Krohn was later
confirmed by Kovalevsky and Mechnikov. In this work also,
Krohn reported one important discovery: "On the contrary to
Medusa aurita and other above-named medusae, "50 he wrote,
"Pelagia noctiluoa develops without the generation of
helminths" (p. 469). Krohn could trace how the swimming planula Velagia, while not settling on the bottom and not
transformed into scyphistomae, forms on the edges of the
mouth orifice processes, later becoming part of Ephyra
(Figure 43, B and C) with marginal sensory bodies. The
citation of this discovery can be found either in later
investigators of the embryology of medusae (for example,
KovalevskySl and Mechnikov (134)], or in textbooks. 52
49. Cited in I. I. Mechnikov, IZBRANNYE BIOLOGICHESKIE
PROIZVEDENIYA (1950), pp. 271 - 472.
50. Krohn compared the development of Pelagia with the
development of Medusa, Cyanea, Chrysaora, Cephea, and
Cassiopea .
In 1861, during his residence in Nice, Krohn observed
the reproduction and development of hydromedusa Eleutheria , 53
The ova arise between ecto- and endoderm and there they
develop into the larval stage of larva. Ectoderm, covering
the embryonic chambers, swells into hillocks, which subsequently break and release young. The larvae are considered
typical planulae and are subjected to the same transformation
as in the planula Cladonema.
The budding occurs not only in asexual, but also in
completely differentiated bisexually related individuals.
Krohn described the process of budding, and noticed that the
budding begins in very young individuals, which are still not
completely separated from the maternal individual.
51. A. O. Kovalevsky, "Observations on the development
of Coelenterata," IZV. OBSHCH . LYUBIT. ESTESTV.
ANTROP. I ETNOGRAFII, 10 (1874), vyp. 2, pp. 1 pp. 1 - 36. To the work of Krohn there is reference
on p. 7. ^
52. K. N. Davydov, TRAITE D 'EMBRYOLOGIE COMPARES
DES INVERTEBRES (1928), p. 78. The drawings given
by Davydov (Figure 36) illustrating the development
of Velagia were taken by him from the work of Delap,
published more than fifty years after Krohn' s. They
are not a bit better than Krohn ' s drawings .
53. A. Krohn, "Beobachtungen iiber den Bau und die
Fortpf lanzung der Eleutheria Quatref. , " ARCH .
NATURG., 27 (1861), 1, pp. 157 - 170.
Figure 43. Later stages of development of medusa Pelagia
noot-iluca (by Krohn) .
a — anterior; b— -posterior end of the body;
stomach.
The development of worms54 Krohn described in many
separate reports. In 1851 he wrote an article on the reproduction and larval stages of "gefirei," 55 He established
the fact of dioecious Phasoolosoma and described the structure
of mature ova of Sipunculus nudus . Attempts at artificial
insemination of these ova were unsuccessful, and Krohn had
to be satisfied with the study of the larvae of Sipunculus
caught in plankton, whose description constituted the final
part of the work. Two small remarks by Krohn are concerned
with the vegetative reproduction of the annulated worm Syllis
and Autolytus . 56 He found in them this change of sexual and
asexual reproduction, which permitted comparison with true
alternation of generations. Many years later 57 Krohn again
turned to the study of reproduction in sillids, describing
the new viviparous species of polychaeta of this genus.
54 . Of the types of worms known in the mid- nineteenth century, many were distinguished later on in independent
groups of forms, including phoronids and chaetognaths ,
and are so listed.
Other of Krohn 's reports separate information about the
development of nemertineans, phronids, and chaetognaths .
The description of larvae and partial transformation of the
first two forms constitute the contents of a special
article^ (135) . On the question of the development of
nemertineans inside pilidium, Krohn inclined to the opinion
that pilidium is considered a helminth, giving origin to
worm-shaped sexual generation. Actinotrocha, in Krohn' s
opinion, is a larval stage of any worm, tentatively relating
to echiuroids. The process itself of the transformation of
actinotrocha he did not observe and noted only the disappearance of the larval organs and the concentration of antennae
in the circumoral corona.
In his excellent investigations on the structure of
chaetognaths, Krohn added the study of their development.
55. A. Krohn, "Uber die Larve des Sipunoulus nudus nebst
vorausgeschikten Bemerkungen liber die Sexualverhaltnisse
der Sipunculiden," ARCH. ANAT . , PHYSIOL. (1851),
pp. 368 - 379.
56. A. Krohn, "liber die Erscheinungen bei der Fortpflanzung
von Syllis prolifera und Autolytus proKfer," ARCH.
NATURG., 18, 1 (1852), pp. 66 - 76; "Uber die Sprossling
von Autolytus prolifer Gr.," ARCH. ANAT., PHYSIOL.
(1855) , pp. 489 - 490.
57. A. Krohn, "Uber eine lebendiggebarende Syllisart," ARCH.
NATURG., 35 (1869), pp. 197 - 200.
58. A. Krohn, "Uber Pilidium und Actinotrocha," ARCH. ANAT.,
PHYSIOL. (1858), pp. 289 - 301.
In a letter to Johannes Miiller sent on February 2, 1853
from Messina, 59 he reported: "I have at the same time studied
the development of sagitta. What Darwin has said about it
relates to the development of any fish" (p. 141) (136).
Krohn did not publish any further special work on the
development of sagitta.
Of Krohn' s two articles on Cirripedia, one concerns
the structure of the cement gland of Lepas anatifera and
Conohoderma virgata and the anatomy of the female genital
system of Lepas and Balanus trintennabulwn . The other^O
includes some data about larval development. Krohn described
the intermediate stage between the young larva which is
similar to the nauplius of the copepods and the late cirripesshaped larva. The work is illustrated with graphs of very
young larva of cirripeds, and also larvae of Balanus species
and Lepas antifera.
The development of gastropodan molluscs (pteropods and
heteropods) was the subject of four reports, the last with
the character of a detailed monograph. ^1 in these investigations his main attention is given to a detailed description
of larvae of pteropods (subclass opisthobranchia) : Cymbulia
Peronii, Tiedemannia neapolitana, Gastropteron Meokelii 3 and
also the larvae of carinate molluscs (subclass prosobranchia)
Pterotraohea (two species) , Carinaria mediterranean and
Fir-iolides (Figure 44) .
59. ARCH. ANAT., PHYSIOL. (1853), p. 137.
60. A. Krohn, "Beobachtungen iiber die Entwickelung der
Cirripedien," ARCH. NATURG . , 26, 1 (1860), pp. 1 - 8.
61. A. Krohn, "Beobachtungen aus der Entwickelungsgeschichte
der Pteropoden, Heteropoden, und Echinodermen. Brief 1.
Mitt. a. d. Herausgeb.," ARCH. ANAT., PHYSIOL.
(1856), pp. 515 - 522; "Beitrage zur Entwickelungsgeschichte der Pteropoden und Heteropoden," ibid . (1857),
pp. 459 - 468; "Uber die Schale und die Larven des
Gastropteron MeckelM," ARCH. NATURG., 26, 1 (1860),
pp. 64 - 68; BEITRAGE ZUR ENTWICKELUNGSGESCHICHTE
DER PTEROPODEN UND HETEROPODEN (Lepizig, 1860),
46 pp.
Figure 44. A-Larva Ctio from the dorsal side with not
completely straightened fans of sailing;
B^— extracted from ovum larva F-ivioZides ,
view from above (by Krohn) .
The accuracy of Krohn's observations concerning the
structure of the larvae of gastropodan molluscs is noted in
recent books on comparative embryology, For example,
K. N. Davydov wrote that "Already, long ago, zoologists turned
their attention to the development of gastropodan molluscs,
and Krohn and Nordmann left for us memoirs which even at
present do not lose their significance" (p. 625) .62 ^ n( j
elsewhere: "It is known that the classical case of heteropods
(_Firzolid.es) was described by Krohn in 1860" (p. 651).
A significant place in Krohn's scientific heritage is
occupied by his investigations on the development of
echinoderms; they were described in no less than ten special
reports and transitional notes in works describing other
subjects. His systematic study of the embryology of echinoderms
Krohn began in 1848 at the time of his three-month stay in
Nice. In February-April he experimented with the artificial
insemination of the ova of the echinoid Echinus lividus . His
observations were compared with data published shortly before
by Derbes, 63 w ho also studied the development of ichinus
brevispinosus .
The mature ovum of Echinus lividus, according to Krohn,
is covered by membrane (chorion) and composed of yolk (this
term Krohn called the ooplasm with nutritional inclusions),
with an embryonic vesicle (nucleus) and embryonic spot
(nucleolus) . Derbes assertion that the embryonic vesicle
disappears in the mature ova up to fertilization, Krohn
considered the result of Derbes' insufficiently thorough
investigation. Within approximately half an hour after fertilization of the ova, the, membrane is separated from the yolk,
a phenomenon, which Krohn explained by processes of endosmosis
and exosmosis: "the ovum membrane absorbs the fluid from the
surroundings and again gives it, to its internal surface"
(p. 6). After fertilization the embryonic vesicle and the
embryonic spot are no longer seen. In the place of the ovum
nucleus, not far from the surface of the ovum, Krohn saw an
empty vesicle. With Baer and Derbes, he considered this vesicle
the nucleus of the fertilized ovum and suggested that the
process of yolk division can begin only after the appearance
of this nucleus. Within three to four hours after fertilization the division of the nucleus begins. Referring to Baer's
excellent observations, Krohn left out the description of
the initial processes of the division. In the conclusion of
the monograph he noted that the result of the yolk division
is the formation of cells, from which the body of the embryo
is formed, because the globules of division are unnoticeably
transformed into cells of the developing larva. The evidence
of this transformation is found in the contents of the cells;
"numerous molecules inside the last substance are nothing
but yolk granules, from which the division globules are
previously formed" (p. 29). "In the same genetic relation,
the nuclei of the cells are related to the vesicular nuclei
of the division globules" (p. 30).
62. Davydov, TPAITE D ' EMBRYOLOGIE .
63. Derbes, "Observations sur le mecanisme et les
phenomenes qui accompagnent la formation de l'embryon
chez l'Oursin comestible," ANN. SC . NAT., 3 Ser.,
8 (1847) , pp. 80 - 98.
64. a. Krohn, BEITRAG ZUR ENTWICKELUNGSGESCHICHTE DER
SEEIGELLARVEN (Heidelberg, 1849), 36 pp.
Krohn prefaced the characteristic of the fully formed
echinus larva with the development of the larva, suggesting
that knowledge about the final development must help the
understanding of the phenomena leading to it. The external
form of the pluteus, the structure of its skeleton and the
digestive system are represented in the drawings and are
described as follows. The formed larva, within eleven days
after the fertilization, is pear-shaped (Figure 45, F) ; it
possesses two pairs of limbs: the short ee and the long dd.
On the convex side, facing the mouth £, and situated between
the limbs, the anal opening c^ is present, although frequently
closed, which is why Johannes Miiller missed it in the larvae
of ophiuroids and echinus. The bilateral symmetry of the larva
is absolutely clear and is expressed in the pairing of limbs
and the calciferous skeleton carrying their branches, in the
situation of the mouth and the anus in the plane of symmetry.
Inside the larva is a cavity extending to the end of the limbs;
the digestive tract lies in this cavity, surrounded by loose
fibrous tissue. The larval surface is covered with skin; the
last is formed from twinkling cells, in each of which there
is a nucleus with nucleolus. The calciferous skeleton consists
of four pairs of toothed branches (Figure 45, G) , the longest
branches gg_ are club-shaped, the next pair is found in the
long limbs hh, another pair in the short limbs kk, and the
last pair H_ is situated across the longitudinal axis of the
larva.
The digestive canal consists of three parts, the anterior
(the pharynx) , the middle (stomach) and the posterior
(intestines), all covered by cells with cilia, similar to
the cells of the skin. The digestive canal is attached to the
body cavity by fibrous tissue; it forms a network united with
the internal surface of the skin. In the junctions of the
network, nucleus-like formations are situated; they are
numerous also under the skin, in the neighborhood of the
calciferous skeleton. Krohn noted Johannes Miiller's error
when he assumed them to be strong fibers instead of nerves,
and the nuclei of fibrous tissue instead of nerve ganglia.
The larva swims with its limbs, mouth opening forward; its
movement is carried out by the activity of the cutaneous
twinkling cilia, which also drive into the mouth food particles
suspended in the water.
The formation of the larva described here takes place as
follows. After the division is completed, the young spherical
larva rotates in the ovum membranes with the help of the
long cilia which cover all the surface of its body. In the
larva at this time can be observed a closed central cavity
and cover, which can be differentiated, forming the wall of
the latter. Krohn erroneously considered that the wall of
the body consists of many layers of cells. Within a day after
fertilization, when the number of the cover cells becomes
sufficiently large, the ovum membrane is torn and the larva
begin to swim. Soon after hatching it acquires its ovoid form
(Figure 45, A); during swimming its narrow end is directed
forward.
In the body cavity of the larva, from the side of the
blunt end, an accumulation of closely situated dark bodies,
similar to the nuclei, is seen. The number of these nuclei
at first is small, but it quickly increases so that they are
found filling half of the body cavity of the larva, toward
its blunt end (Figure 45, A,e) . Later, nuclei are separated
from each other and distributed evenly in the body cavity, and
then partially accumulate not far from the rudiments of
the calciferous skeleton. After this, the nuclei become angular
or fusiform and begin their transformation into the fiber of
the reticular tissue which strings the body cavity and holds
the digestive tract.
Figure 45. Development of echinus, Echinus liwidus (by Krohn) .
A — larva shortly after hatching: a — blunt pole; b — sharp pole;
c, c — cover; d — central cavity; e — accumulation of "nucleuslike formations" from which the fibrous tissue is later
formed.
B — "ideal sketch" of the larva, already having the form of the
body with three surfaces: a, b, c — anterior, posterior, and
upper surfaces; d — anterior angle; e, e — posterior angles;
f — lower angle ; g — anus .
C— — cross-section of this same stage of development, in which
for elucidation of the passage the previous apple-shaped
form was drawn: a, b, c, d, e, f, g — as in figure B;
h — rudiment of the digestive tract in immature form;
k — its rudiment in later form.
D — larva of the following form, back view: a, b — posterior and
upper surfaces; c — anus (back passage) ; gg, 11, kk — rudiments
of club-shaped, arch-shaped, and transverse branches of
calcareous skeleton.
E — later larva: a, b : c, g, k, 1 — as in figure D; dd — rudiment
of posterior limbs; hh — first traces of hydrants in it.
F — larva of pear-shaped form, back view: dd — posterior limbs;
ee — anterior limbs; f— -mouth; q — -anus; ssss — cilia apparatus.
G — calcareous skeleton of the formed larva, represented in
an isolated view: gg — club-shaped hydrants; hh — hydrants of
anterior limbs; kk — arch-shaped hydrants; 11 — transverse
hydrants .
From Krohn's description it is clear that he saw, for
the first time, in the cavity of echinus division that
accumulation of cells, which was called by the latest
embryologists the primary mesenchyma. He correctly determined
the fate of these cells in larval development. This discovery
of Krohn's was so thoroughly forgotten that Mechnikov, who was
well acquainted with the old embryological literature,
attributed it to Selenka, whose work was published exactly
thirty years later after Krohn's monograph. 65
"The development of Echinus micro tub eraulatus ," Mechnicov
wrote, "was studied by Selenka. The first differentiation of
the embryonic layers begins with the bulging of the lower
cells of the blastoderm. Until the beginning of their
protrusion they form a number of wandering cells, which later
develop into the cutis and are considered mesodermal or
mesenchymal cells. "66
Krohn observed that after nearly thirty hours after
fertilization the following important changes take place in
the larva of Echinus lividus . In the center of the blunt pole
a small hole-like deepening appears. The blunt pole widens and
thickens; the hole in its center becomes deeper and wider,
so that the body of the larva becomes similar in form of an
apple. The more enlarged hole "formed as a result of that,"
Krohn wrote , caused the skin in this area gradually to protrude
(EINSACKT ODER EINWARTSSTULPT) into the cavity
of the body. The protrusion (EINSACKUNG) submerged
deeper in the body cavity and extended into a canal
which finally reached the walls of the body cavity
toward the blunt pole. The sac which appeared by this
means stretched through the body cavity (Figure 45,
C, h, k3 and is the rudiment of the digestive cavity.
The edge or circumference of the primary hole becomes
a leading opening in the canal, which is the anus. In this view's favor — on the manner of appearance of
the digestive tract — is the fact that it also
indicates that the wall of the canal is absolutely
equivalent to the skin in thickness and structure.
(p. 18)
65. E. Selenka, "Keimblatter und Organanlage bei Echinicen,"
ZTS. WISS. ZOOL.,, 33 (1879) .
66. I. I. Mechnikov, "Vergleichend-embryologische Studien.
3: Uber die Gastrula einiger Metazoen," ZTS. WISS. ZOOL.,
37 (1882) , pp. 286 - 313.
Krohn's discovery of the nature of the formation of the
larval intestine of echinus undoubtedly possesses outstanding
significance. This, unquestionably, is the first description
of the invaginated gastrulation phenomenon. Its study subsequently played a great role in the progress of comparative
embryology. Comparative analysis of methods of separation of
the endoderm is included in the basis of Kovalevsky f s evolutionary concept about the formation of the embryonic layers,
and also in the basis of Haeckel's gastrula theory. Krohn's
priority in this question was not definitively underlined.
Mechnikov, in the article just cited, wrote the following.
Derbes (1847) described the formation of the larva of
Echinus esoulentus and mentioned a stage in the form of
a double sac with skin layer in which the caecum opened
to the outside. Twenty-five years later, Haeckel gave
this the name gastrula, which was accepted by scientists
all over the world. Derbes thought that the opening
of the rudiment of the intestine was the mouth, but
August Krohn (1849) showed that it corresponded to the
anus of the pluteus. He described the process of
protrusion itself.
Krohn referred to Derbes' observations: "The reader can form
an excellent concept of the gradual formation of the digestive
tract just described by looking at Figures 13 and 14 in
Derbes' article; however, the author apparently did not pay
attention to the process of formation itself" (p. 19). The
following is written by Derbes: "The spherical form of the
larva is changed by pressure at one point of the surface.
Gradually this pressure becomes more pronounced, and its
center is penetrated by an opening which leads to a rudiment
of the intestinal cavity. Beginning from this moment, the
movement of this opening is always directed forward and, later,
upwards. . . that is, the mouth looks towards the zenith"
(pp. 91 - 92). It is clear that Derbes did not put the deepening
on the surface of the spherical larva in genetic relation with
the formed intestine. According to his opinion, the opening
formed in the center of the deepening united in an unknown way the developed rudiment of the digestive tract with the
external world. Krohn pointed also to DerbSs' mistaken assumption that this opening was the mouth. He established that the
opening in the area of protrusion is the anal opening, and
the mouth is formed in another place significantly later
(on the fourth day after fertilization] .
Further observations of Krohn are concerned with the
changes in the larval form" as it becomes bilaterally symmetrical (Figure 45, B and C) . He described the formation of
limbs and the calciferous skeleton (Tigure 45, D and E) ,
which gradually acquires a different configuration.
In the following year, Krohn repeatedly returned to the
study of development of the different echinoderms and
published many reports on this subject, One of these reports
considered the development of the holothurian and echinus. 67
The larvae of holothurians (Holothuria tubulosa} were
obtained from plankton, because attempts at artificial
insemination proved unsuccessful. The youngest larva observed
by Krohn "is similar to an elongated egg (Figure 46, A); in
its sharp pole there is an opening leading to a sac-like
protrusion in the body and an ampul la- shaped canal widening at
the end Ik This sac is the rudiment of the digestive tract,
the opening doubtlessly is the anus" (p. 345). Krohn noticed
that the larva of Holothuria tubulosa, at this stage, is very
similar to the larva of Echinus lividus described earlier by
him (1849) . Similar to the latter, the larva of Holothuria
tubulosa is covered by cilia, and with their help it swims with
the imperforate pole forward. The surface of the body and the
digestive tract consist of cells which, with their nuclei,
become noticeable with the addition of fresh water. In the
body cavity, as in the larvae of Echinus lividus, fibrous
tissue with fusiform cells is found. Later (Figure 46, B) on
the abdominal side of the larva a depression appears — which
is the future transverse fissure of ausicularia; in this
fissure the mouth opening later appears. Later the body
acquires a kidney-shaped configuration (Figure 46, C) . The
digestive tract, at this time, forms the rudiments of the three
parts, pharynx e, stomach d and intestines c. The appearance
of the mouth opening coincides with the beginning formation
of the ciliary strings in the auricularia. At this stage there
is some data about the transformation of the Echinus lividus .
67. A. Krohn, "Beobachtungen aus der Entwickelungsgeschichte
der Holothurien und Seeigel," ARCH. ANAT . , PHYSIOL.
(1851) , pp. 344 - 352.
Krohn's other reports on the development of echinoderms
briefly designate some representatives of the type68 or
species. 69
Special attention must be given to Krohn's investigations on the development of tunicates. Originally his interest
was attracted by Salpa, in which the wonderful phenomenon of
the alternation of sexual and asexual generations had already
been recognized. This discovery belongs to the poet Adalbert
Chamisso (1781 - 1838) . With the Dorpat-born zoologist
J. F. Eschscholtz, he traveled around the world on the
Russian ship RURICK. Chamisso published the results of his
investigations in "On Some Animals of the Linnean Class of
Worms, noted during a world tour, performed by Count
N. Romanzoff, under the command of Otto von Kotzebue, from
1815 to 1818, Part I: On Salpa. "70
68. A. Krohn, "Bemerkungen uber einige Echinodermenlarven, "
ARCH. ANAT., PHYSIOL. (1851), pp. 353 - 357; "Uber
die Entwickelung der Seesterne und Holothurien (Brief 1.
Mitt. a. d. Herausgeb . ) , " ibid . (1853), pp. 317 - 321;
"Beobachtungen iiber Echinodermenlarven (Brief 1. Mitt,
a. d. Herausgeb.)," ibid . (1854), pp. 208 - 213; "Uber
neuen Entwickelungsmodus der Ophiuren," ibid . (1857),
pp. 369 - 375.
69. A Krohn, "Uber die Entwickelung einer lebendiggebarenden
Ophiure (Brief 1. Mitt. a. d. Herausgeb.)," ARCH. ANAT.
PHYSIOL. (1851), pp. 338 - 343; "Uber die Larve von
Spatangus purpureus (Brief 1. Mitt. a. d. Herausgeb.),"
ibid . (1853) , pp. 255 - 259; "Uber die Larve des Echinus
brevispinosus ," ibid . (1853), pp. 361 - 364.
70. DE ANIMALIBUS E CLASSE VERMIUM LINNEANA IN
CIRCUMNAVIGATIONE TERRAE AUSPICANTE COMITE
N. ROMANZOFF DUCE 0TT0NE DE KOTZEBUE ANNIS
1815, 1816, 1817, 1818 PERACTA OBSERVATIS
ADELBERTUS DE CHAMISSO, Fasc. primus: DE SALPA
(Berlin, 1819) , iv + 24 pp.
Figure 46. Three stages of the development of the sea
cucumber Holothuria tubulosa (by Krohn) .
A and B: a — anus; b — digestive cavity,
Id — anus; c, d, e — rudiments of the
:estine, stomach and pharynx.
C:
inte
The alternation of sexual and asexual generations in
Salpa was described by Chamisso as follows:
The species of Salpa is found in double form: each
generation of the species is dissimilar to its
parents, but through birth posterity is similar to the
last, so that any Salpa differs from its parent but
is identical with its grandparents. Both forms are
similar to headless molluscs, hermaphrodites or the
female sex. Both of them are viviparous, but one of
them is a solitary animal, the originator of many
descendants . The other represents a complicated
branch consisting of animals, each united with the
others by the necessary connection which gives birth
to one descendant. These changed forms of the unchanged
species are called solitary (Proles solitaria) and
aggregated or colonial (Proles gregata) generations,
(p. 2)
After the discovery by M. Sars^l of the analogical
change of generations in scyphomedusa, Steenstrup united
these facts to produce one biological regularity. Steenstrup 's general conclusion was highly regarded by his
contemporaries, in particular by Baer.72 Krohn also gave
great significance to Chamisso's discovery and Steenstrup 's
ideas. He made it his task to study in detail the reproduction
and development of Salpa. To do this, he settled for many
months on the coast of Sicily, where the sea provided him the
necessary material. Krohn put the results of his observations
into a special work, 73 in which, first of all, he completely
confirmed Chamisso's observations, by distributing them over
seven species of Salpa which were, for the first time, partly
described by him. The comparison of the solitary and colonial
forms (Proles solitaria and Proles gregata, in Chamisso's
terminology) allowed Krohn to regulate the taxonomy of this
group of tunicates. He showed that salpae described under
different names frequently proved to be different stages of
the development of one and the same species (137) . Later,
Krohn gave the characteristics of the structure of the
heteromorphic generation, and also described the ovum, the
seminal glands, and the process of fertilization in the
sexual generation.
Within the present book, the greatest significance is
placed on the section in Krohn' s article (111) in which he
discussed development of the embryo in the maternal organism.
After fertilization, the embryonic vesicle and the embryonic
spot disappear, after which the ovum enlarges in size and
acquires a regular spherical form. This was according to Krohn,
who was not completely convinced of its authenticity. Sometimes
the ovum is not seen like that, and in its place a round body
appears, raising a region of tunica of the mother and jutting
into its cavity in the form of papilla. This body, Krohn wrote,
is nothing other than the rudiment of the placenta which, by
deepening in the cavity of the body of the maternal organism,
enters in connection with two of its blood vessels. Only
after the formation of the placenta does the embryo begin to
develop, at first in the form of a very small body appearing on the summit of the placenta under its cover (138) . In this rudiment of the embryo all organs of the last are developed;
"however," Krohn wrote, "all that concerned its development
during the first period remained for me almost completely
unknown" (p. 123). He could only establish that one of the
first organs of the embryo by the time of appearance is "the
respiratory cavity." The embryo changes from compact to hollow,
following which the rudiments of the branchae and nervous
ganglion are already seen, while the organs "visceral nucleus"
and heart become noticeable only later. Only after that does
the embryo acquire a definite form; the anterior and posterior
openings appear in it. At the end the embryo becomes more
voluminous than the placenta, and all its organs intensively
enlarge, especially the nervous ganglion, from which numerous
nervous branches grow. At the same time muscular strips and
blood vessels appear, which are not completely formed.
71. M. Sars, "Uber die Entwickelung der Medusa aurita und Cyanea
caprtlata," ARCH. NATURG . , 7 (1841), pp. 9 - 34.
72. See Chapter 23.
73. A. Krohn, "Observations sur la generation et le developperaent des Biphores (Salpa)," ANN. SC . NAT., 3S6r.,
Zool., 6 (1846), pp. 110 - 131.
Krohn' s concluding paragraph of the work is dedicated
to the processes of budding in the asexual regeneration of
Salpa, and to the formation of colonies; the character of the
last varies in different species. Here the description of the
stolon and the embryos is given, situated along it so that
their axes cross the stolon at a right angle. These embryos
develop in definite succession, depending on their situation
on the stolon.
The development of salpa, especially the formation of
their embryos from fertilized ova, represents one of the most
difficult principles of embryology. Krohn' s investigations
began this study and recent opinions are credited to many
Russian embryologists at the end of the nineteenth and the
beginning of the twentieth century, including A. 0. Kovalevsky,
M. M. Davydov, A. A. Korotnev, and V. V. Zalensky. 74
Six years after the publication of the above-mentioned
work, Krohn published an article presenting the results of
his investigations of the little-studied group of tunicates,
the doliolum.75 Q U oy and Gaimard made a voyage on the
74. "The first investigations of the development of Salpa go
back to. Krohn," K. N. Davydov wrote in his handbook
CTRAITE, p. 867) .
75. A. Krohn, "Uber die Gattung Doliolum und ihre Arten,"
ARCHIV. NATURG., 18 (1852), pp. 53 - 65.
ASTROLABE 7 ^ to Road Island, Ambon (in the Moluccas), and
to the coast of the Vanikoro Islands (between the New Hebrides
and the Solomon Islands) . During this time they discovered
this small transparent tunicate, which they described and
presented only incompletely, and for which they suggested the
generic name Doliolwn.
Later T. Huxley, on a tour around the world aboard the
ship RATTLESNAKE, saw doliola in the southern part of
the Pacific Ocean and described their structure more exactly
and in more detail than Quoy and Gaimard,77 but in his notes
the male genital glands were given for individuals of sexual
regeneration.
Krohn, for the first time, discovered the presence of
doliola in the Mediterranean Sea (near Messina and Naples) ,
studied the structure and described three new species.
According to Krohn, doliola "are free living ascidians, but
in many respects are similar to salpa and form, therefore, an
interesting intermediate link between both these orders of
tunicates" (p. 53). From Krohn's description, the structure and
reproduction of doliolum, according to the majority of the
features, must be related to salpa; this applies also in the
present taxonomy.
Krohn observed that from the ova of doliola cercarialike larvae develop which later undergo metamorphosis .
The peculiarities of the larvae are connected, according
to Krohn, with the mode of life of the tunicates more than to
ascidians, which are in their adult condition fastened motionless to the substrate. In accordance with this, in the ascidian
larvae the tail disappears early, but in the larvae of doliola
it remains throughout metamorphosis and serves as an organ
of movement. The process of reduction of the tail in the larvae of doliola takes place as in the larvae of ascidians . In
the latter the disappearance of the tail was noticed for the
first time by Milne-Edwards in an example of the colonial
ascidian Amouruoium proliferum, and was described in more
detail by Krohn for Phallus-La mammillata .
76. VOYAGE DE DECOUVERTE DE L' ASTROLABE EXECUTE PAR
ORDRE DU ROI PENDANT LES ANNEES 1826 - 29 SOUS
LE COMMANDEMENT DE M. J. DUMONT D'URVILLE.
ZOOLOGIE PAR MM QUOY ET GAIMARD, vol. 3 (1834).
77. T. Huxley, "Remarks upon Appendicularia and Doliolum,
Two Genera of Tunicates," PHIL. TRANS. ROY. SOC .
LONDON (1851), pp. 599 - 602.
The metamorphosis of doliola was again studied by Krohn
on the species Doliolum Nordmanni. Krohn represented two
stages of transformation; Figure 47, A represents the stage
when the tail of the larva has not begun reduction; and
Figure 47, B, the larva with shortened tail. Until metamorphosis the tail of the larva is tapered, with the two ends
covered with gelatinous membrane a_; its axis d_ is composed
of cuboidal cells situated in one row, so that it seems to
be segmented. The root of this axial shaft penetrates into
"the vesicular appendix" c_, lying under the digestive tract
of the larva. This vesicle decreased simultaneously with the
reduction of the tail and at the end it disappears (139) . In
the stage represented in Figure 47, B, all organs in the
larva are already differentiated; on its dorsal side there is
the placing of the stolon £.
Krohn ! s description of the structure of the larva of
doliolum and its transformation distinctly shows that he saw
in it all the principal features of organization, on the basis
of which the doliolum was later counted as a type of chordate
(subtype tunicate, class salpa) : the presence of the reduced
cord at the time of metamorphosis ("axis of tail"), the situation of the nervous ganglion o_ on the dorsal side and the
heart m on the ventral side.
Correct evaluation of Krohn ! s investigations was given
by V. N. Ulianin in the classic monograph on doliola, first
published in Russian, and two years later in German.' 7 ** Ulianin
wrote the following: "Soon after the appearance of Huxley's
article the doliolum was found in the Mediterranean by Krohn.
78. V. Ulianin, ON THE DEVELOPMENT AND REPRODUCTION OF
DOLIOLUM (Moscow, 1882), 100 pp. B. Uljanin, DIE ARTEN
DER GATTUNG DOLIOLUM IM GOLFE VON NEAPEL UND DEN
ANGRENZENDEN MEERABSCHNITTEN, FAUNA UND FLORA
D. GOLFES V. NEAPEL, X. Monographie (1884), 140 pp.
The article in which this discovery was published possesses
important significance in the history of doliolum, as it
contains the first observations on the development of this
animal. Krohn not only described the tailed larva of doliolum,
tut he also pointed out the alternation of generations in the
reproduction of this animal.... On the basis of his observations Krohn concluded that from the ovum of doliolum is formed
the tailed, freely swimming larva, which after the
loss of the tail is transformed into an asexual doliolum"
(p. 2). Later on Ulianin noticed Krohn's mistake in dividing
doliolum according to the number of muscular strips. In
all species of sexual generation there are eight, but in
asexual species there are nine strips. Therefore the species
described by Krohn, Doliolum Trosoheli , is in fact an asexual
individual D. dentioulatum Q. and G., and D. Nordmannz Krohn
is the asexual generation of D. Mulleri Krohn. Particular
significance was given by Ulianin to Krohn's embryological
observations. "All that is known presently about the
embryological development of Doliolum," Ulianin wrote,
"comes exclusively from Krohn, who for the first time described
the free larva. All later authors.... only redescribed it, not
adding anything essential to Krohn's description (p. 47).
The central place among Krohn's investigations of the
development of tunicates is occupied by his work concerning
the solitary ascidians.? 9 The artificial insemination in
ascidians which was used successfully for the first time by
Baer, Krohn also used, observing the development of Phallusia
mammillata step by step for three months. He described the
mature ovum of this ascidian in the following way. The ovum
present in the ovum-fluid is supplied by papillae and covered
by a cover membrane under which the proper ovum membrane is
present. Somewhat deeper lies a hyaline membrane containing
inclusions which is green in color. The yolk itself is
colorless, the embryonic vesicle and the embryonic spot in
the mature ova are unnoticeable . The above mentioned green
hyaline membrane was considered by Krohn, following MilneEdwards, a source of formation of tunica. The error of this
view was later established, but it was repeated in many
79. A. Krohn, "fiber die Entwickelung der Ascidien," ARCH.
ANAT., PHYSIOL. (1852), pp. 312 - 333. In the following
year this work was published in English as "On the
Development of the Ascidians," SC. MEM. NAT. HIST.
(1853) , pp. 312 - 329.
subsequent embryological works . The division of the ovum
begins two to three hours after contact with the sperm.
Krohn considered that the division, at least in the first
stages, follows the rule of progress. The vesicular nuclei
of the globules of division disappear before every division
and then again become visible. "Instead of nuclei," Krohn
wrote,
in every divided globule an absolutely peculiar
distribution of yolk molecules is noticed. Namely,
they are distributed in the form of strands, which are
directed from the depth, from the medial point by
radius in all directions to the lighter periphery of
the ovum and, apparently, come out from two centers of
irradiation. After the end of division, inside the
new globules the nuclei again become noticeable, then
these radiant figures disappear and the yolk granules
are found to be situated close to each other, (p. 315)
These observations show that Krohn exactly described
many details of mitotic division in blastomeres.
Embryonic development goes quickly, and a day after
fertilization a cercaria-like embryo with a more or less
developed tail is already present in the ovum membrane. The
body and the tail of the embryo are composed of cells which
are especially noticeable on the surface. The cells have a
polygonal form, and contain granules and nuclei in the center.
The axis of the tail, according to Krohn 1 s description, is
composed of larger rectangular cells with nuclei situated in
a row, one following the other, and therefore they have a
striated or a disjointed form (Figure 48, A, b) .
Shortly before the final formation of the larva the tail
undergoes remarkable transformations. According to Krohn' s
observations, they amount to the following. The axis is
transformed into a canal, as its cellular structure gradually
disappears due to the destruction of the partitions between
the neighboring cells and the liquefaction of their contents.
The small cells surrounding the central tail strand are
transformed into longitudinal muscular fibers. On the dorsal
side of the larva at first appears one, and then behind it
another, pigmented spot of granular origin (Figure 48, A, d, e
and B, e, f ) . At the time of transformation this formation is destroyed, and the pigment passes into the blood channel.
The formed larva is set free from the membrane by the tail
movement. The body of the larva CFigure 48, B) in the anterior
end is supplied by three similar processes on the sucker. The
larva is soon attached at the anterior end and undergoes
transformation, one of its marks being the disappearance
of the tail. Milne-Edwards saw only that the axial part of
the tail is set free from its covered sheath and extends into
the body of the larva, but he did not elucidate the subsequent
fate of this formation. "By my observations," Krohn noted,
the setting free and the extension of the tail axis ,
the deep immersion by the tail in the body of the larva
only precedes the processes of reduction which it soon
undergoes. Directly after the extension, the tail axis
remains undamaged at the posterior part of the body.
It is situated here convoluted into a spiral coil . . . .
With the beginning of the development of the young
ascidian, this coil first disintegrates into a large
number of strips situated close to each other, which
then are gradually destroyed; the number and size of
the strips decrease, but the insignificant remnant
does not disappear entirely. (Figure 48, C) (pp. 318 319)
Krohn himself considered his observations on the development of ascidians incomplete, and he acknowledged only the most
essential changes. He described in particular the formation of
the vessels of the tunica and the development of the respiratory cavity, or gill cleft, and behind it the rudiment of the
digestive canal in the form of a loop-shaped canal . Somewhat
later three openings on the spinal side of the body appear: the
most anterior, the inlet into the respiratory cavity and
digestive canal, and two posterior which later merge together
in a common excretory opening. Simultaneously, the nerve
ganglia develop in an elongated formation in the middle of the
back near both pigmentation spots. Near the nerve ganglia the
rudiment of the muscular strands form and the dorsal fissure
appears. The digestive canal is differentiated into three
parts: a canal which opens into the respiratory cavity, stomach,
and intestine. In the walls of the respiratory cavity there
develop near the stomach the first branchial clefts with cilia
at the edges, and at the ventral fissure the heart develops,
possessing the form of a short duct. The metamorphosis is
completed by the specialization of the gill-clefts and the
development of siphons .
Figure 47. Larvae of Doliolum Nordmanni .
A — Larva up to transformation: a — larval
membrane; b— young doliolum; e, f — posterior
and anterior opening; d — axis of the tail;
B — larva of doliolum in the beginning of
transformation: a, d, e, f — as in A;
c — "vesicular appendix; g— wall of the
respiratory cavity; h — -digestive tract;
k — stomach; 1 — intestine; m — heart; n— ventral
fissure; o — nervous ganglion with outgoing nerves;
p—— the third from back muscular strand, penetrating the rudiment of the stolon (q) (by Krohn) .
Krohn's work represents the first systematic description
of ascidian development in world literature; it remains incomplete and not free from mistakes, which, of course, does not
reduce the historical significance of this undoubtedlyremarkable investigation. But the exact and detailed study
of the embryology of ascidians belongs to A. 0. Kovalevsky.
In his work, their relationship to the vertebrates was proved,
delivering a fatal blow to the metaphysical theory of types
in the animal kingdom. It formed the basis of comparative
evolutionary embryology, first advanced by Krohn. In the work
dedicated to the development of ascidians, 80 Kovalevsky wrote
the following:
Leaving aside the investigations of earlier authors,
whose results either are already completely reworked
by present scientists, or, to a lesser extent, are
partially expanded, we must mention Milne-Edwards,
Van Beneden, Kolliker, and, in particular, Krohn. Of
all these investigations the results of Krohn's
investigations are in closest agreement with our own
results. Although he described the accumulation of
pigments, which completely coincides with our observations , he did not discover the walls of the saccule in
which these organs of sensation are situated, and
generally he traced the development step by step. The
formation of the axial strand in the tail of the
ascidian larva was observed by Krohn, although he
explained it as a formation of emptiness in the cells .
Although the transformation of the larva into the
sessile form was described by him in detail, he had
only a slight understanding of the anatomy of the larva
and therefore he could not observe the particular
features, (p. 41)
80. A. O. Kovalevsky, "Istoriya razvitiga prostykh astsidii"
(The history of development of the simple ascidia) (1886) ,
SELECTED WORKS (Izd. AN SSSR, 1951) , pp. 41 - 78.
81. A. Krohn, "Uber die fruheste Bildung der Botryllusstocke,"
ARCH. NATURG., 35 (1869), pp. 190 - 196; Uber die
Fortpflanzungsverhaltnisse bei den Botrylliden, " ibid .,
pp. 326 - 333.
At the end of the 1860s, in Naples, Krohn studied budding
in the complex ascidian Botryllus , and he presented the results
of his observations in two reports. 81 In the first of these articles Krohn disproved the erroneous data of Milne-Edwards
and Sars and confirmed the observations of Mechnikov that the
larva of Botryllus possesses the same simple structure as
that of the solitary ascidian and undergoes analogical
metamorphosis. After settling on the bottom, the young
Botryllus , already in the process of transformation, produces
a bud from which a second individual originates, which in
turn begins to bud. As a result a stellate colony is obtained.
The budding of the colonial ascidian is represented in the
second article, in which Krohn compared its details with the
corresponding phenomena in salpa.
With these fragmentary investigations of vegetative
multiplication, Krohn' s scientific activity apparently came
to an end. In the following years (1870 to 1880) his reports
were regularly placed in journals, but no works appeared after
this time.
For thirty-five years he collected facts from the field
of anatomy and embryology, mainly of invertebrates, covering a
very great number of systematic groups CI 40) . His embryological
works (including descriptions of larvae and means of reproduction) concerned coelenterates, nemerteans, annelids, molluscs,
crustaceans, echinoderms, and tunicates.
Krohn did not belong to those investigators paving new
roads in science. All his comparative-anatomical and comparativeerabryological remarks are concerned with the comparison of
closely related forms. Comparing, for example, representatives
of different classes of echinoderms and establishing the
features of similarity and difference between salpa and
ascidians, Krohn did not offer sympathy either to the theory
of types, nor to the idea of unity of planes, nor to evolutionary study. Krohn cited Darwin only in his works on the
structure and development of cirripedes, highly rating his
monograph dedicated to this order of crustacean.
Besides this, evidence exists about Krohn' s deep interest
in the investigations of A. 0. Kovalevsky, who established the
similarity of the embryonic development of ascidians and
vertebrates. The first report by Kovalevsky dedicated to the
development of ascidians was published in 1866 in ZAPISKAKH
PETERBURGSKOI AKADEMII NAUK (Notes of the Petersburg
Figure 48. The development of the ascidian Phallusia
mammillata (by Krohn) .
A — late embryo with two pigmentation spots: a — body;
b — beginning of the tail; c — rudiment of upper sucking
processes? d, e — anterior and posterior pigmentation
spots .
B — larva, side view: a— tunica with green bodies; bb— the
axis of the tail; cc — its canal; d — horizontal half;
e, f — anterior and posterior pigmentation spots;
g — right anterior; h — posterior sucking processes.
C — Phallusia in the process of metamorphosis: a— widely
opened respiratory siphon; bb— posterior (constrictor)
siphons; c — nervous ganglion with the nervous branches;
d: — digestive tract; e — stomach; f — intestine; g — situating
coil of the larval tail; hhhh — first two pairs of gill
openings of the respiratory sac; i — pigmentation mass over
the nervous ganglion; k — ventral fissure; 1, 1 — tunica.
Academy of Science); two years later an article followed in
NACHRICHTEN VON DER GESELLSCHAFT ZU GOTTINGEN. The
data mentioned there, apparently, did not convince Krohn. In
1871 in ARCHIV FUR MIKROSKOPISCHE ANATOMIE, edited by
Max Schultze, a new work by Kovalevsky was published, 82 a t
which time Schultze added the following postscript to a letter
directed to Kovalevsky on January 18,1871: "Krohn, who read
your article in proof and made minor corrections, sends you
heartfelt regards. At first he was, as you can imagine, very
much against the relationship with vertebrates; 83 b u t then he
began to hesitate." 84
Krohn was one of the pioneers of zoological investigations on the Mediterranean coast, which later became a place
of pilgrimage for naturalists from different countries of
Europe.
It is highly probable that Kovalevsky and Mechnikov were
well acquainted with the works of Krohn. His remarkable
comparative embryological investigations, of which they became
aware in the mid-1860s, revealed the nature of Krohn 1 s
scientific activity and produced an impression on them. Like
Krohn, they spent many years of their lives as travelling
naturalists, more than once following him to those localities
82. A. O. Kovalevsky, "Entwickelungsgeschichte der einfachen
Ascidien," MEM. AC. SC . ST. PETERSB . , VII Ser., 10,
No. 15 (1866), 16 pp.? "Beitrag zur Entwickelungsgeschichte
der Tunicaten," NACHRICHTEN VON DER GESELLSCHAFT
ZU GOTTINGEN, No. 19 (1868), pp. 401 - 415? "weitere
Studien iiber die Entwickelung der einfachen Ascidien,"
ARCH. MIKR. ANAT., 7 (1871), pp. 101 - 130.
The Russian translation of the first and third articles
are to be found in A. O. Kovalevsky, SELECTED WORKS,
editor and commentator A. D. Nekrasov and N. M. Artemov
(Izd. AN SSSR, 1951), pp. 41 - 78 and 79 - 122.
83. The paper was on the relationship of ascidians to vertebrates .
84. Schultze' s letter is included in the book PEREPISKA
A. 0. I V. 0. KOVALEVSKY (Postscripts of A. O. and
V. O. Kovalevsky) , edited by A. A. Borisiak and
S. Ya. Streich. The extract of this letter is published
here with the permission of S. Ya. Streich.
rich in the zoological material of the sea — Naples, Messina,
Nice, Madeira — and using, in particular, those subjects on
which Krohn made many important observations leading to
serious theoretical meditation. Kovalevsky later cited with
respect the works of Krohn on the structure of sagitta and
the development of tunicates, and Mechnikov cited his works
on the development of coelenterates and echinoderms .
In the preparation of that revolution in embryology
which was accomplished by Kovalevsky and Mechnikov, converting
the comparative-descriptive embryology into comparative
evolutionary embryology, Krohn 's modest investigations played
their role, and therefore his name must not be forgotten in
the history of Russian science.




==Comments==
==Comments==


(1) Needham arbitrarily limited the contents of his  
(1) Needham arbitrarily limited the contents of his book absolutely to the history of chemical embryology. In addition, his extreme unobjectivity drew attention, as he dwelt mainly on the works of English authors, while, according to his opinion, the book should have represented the history of embryology of all times and peoples. In the chronological table of scientists of the world embryology, covering up to the beginning of the 20th century (p. 266 of Russian translation) , Needham did not find places for the names of N. A. Warnek, A. 0. Kovalevsky, I. I. Mechnikov and their numerous Russian followers in contrast to
book absolutely to the history of chemical embryology. In  
addition, his extreme unobjectivity drew attention, as he  
dwelt mainly on the works of English authors, while, according to his opinion, the book should have represented the  
history of embryology of all times and peoples. In the  
chronological table of scientists of the world embryology,  
covering up to the beginning of the 20th century (p. 266 of  
Russian translation) , Needham did not find places for the  
names of N. A. Warnek, A. 0. Kovalevsky, I. I. Mechnikov  
and their numerous Russian followers in contrast to  
 
Fr. Balfur, who published more than 70 years ago the first
(and for that time excellent) manual book on comparative
embryology, in which the Russian works were given proper
places (14) .
 
(2) Samples of these "compositions" are pictured on
the prints of that time, reprinted in the book "Kunstkamera
Peterbergskoi Akademii Nauk", 1853, 293 p. (see Fig. 6.10
and 11 of the mentioned book) . Description of anatomical
and embryological "compositions of Ryuish and texts of the
poetical Latin inscriptions which accompanied them much
earlier were mentioned by Baer in "Memories about anatomical
cabinet" ("Memoire uber das Anatomische Kabinet, gelesen in
den Sitzungen der Phys.-math. Kl. d, Sep. 20, 4 u. Oct. 18,
1850; Collection the museum of anthropology and ethnography
in Imp. Ac. Sc, 1900, p. 111-152) (21).
 
 
 
Here and below figures in round brackets show the number
of text page, to which the comment is related.
 
 
(3) Doctor van der Hulst, (see below for data about
him) on April 14, 1724 ,; sent organs which belonged to a
girl, who died of smallpox, in Kunstkamera. Caesar
honored him due to the hermaphrodite (Materials for history
of Imp. Ac. Sc, V. 1 Spb, 1855, p. 38) in the other place
of these "Materials" wrote due to bringing to the Academy
of Science a monster, who was born with two heads, determined by Doctor Stepan Gaokantsk, ...8 roubles were given"
(V. II, on March 9, 1732, p. 119) (23).
 
(4) In "Materials for history of Imp. Ac. Sc." there
are many documents about the obtaining of teratological
objects into Kunstkamera.
 
A letter from Vyborkh by colonel general Karnov says
"his major Beshentsev Fedor Fedrov in the Vyborgskaya
office found a monster of a lamb having four legs and one
neck, and the head had the view of a double one. This head
had two mouths with tongues and two eyes: they were in the
 
middle where the forehead must have been " ('Materials..
 
V. I, Spb., 1855 No. 165, March 9, 1725, pp. 9-97).
 
Inventory of objects, obtained on March 8, 1725
"Number 1: lamb, with 8 legs, another with three eyes,
two trunks, 6 limbs were sent from Tobol'sk by Kozlovsky.
Number 2: baby, with 3 legs: from Lower Novgorod from
Governor Rzhevsky. Number 3: calf with 2 deformed legs:
from Ufa from commandant Bakhmetov. Number 4: baby with
two heads also from Bakhmetov. Number 5: one baby, with
eyes under the nose and ears under neck: from Nezhin.
Number 6 - two babies - breasts and abdomens were joined from Akhtyrok from prince Mikhail Golitsyn; hands, legs
and head were normal. Number 7 - baby, with a fish tail,
born in Moscow in Tverskaya. Number 8: two puppies,
born from a 60-year-old woman, from Akhtyrok from prince
Golitsyn. Number 9: baby with 2 heads, 4 hands, 3 legs:
from Ufa from commandant Bakhmetov" ("Materials...",
V. I, No. 193, March 18, 1625, p. 99).
 
Grigorii Ivanov found a monstrous head puppy with
8 legs ("Materials, V. I, No. 291, September 21, 1725,
p. 145).
 
 
"....Dead body of an infant, with 6 fingers, horn by
a daughter of Tikhanov who was working on a marine ship
Astrakhan. . .given in Kunstkamera" ("Materials..." V. II,
1886, No. 99, November 29, 1731, p. 83).
 
"On January 17 of this same year, my brother sent a
monster which had 2 mouths, the daughter of Ulita Kiryanova
the wife of sergeant Nikivor Kosharov gave birth to it:
this monster had four hands, four legs and two faces, one
had normal view and the other had one eye" ("Materials...",
V. Ill, 1886, No. 13, January 20, 1736, p. 14).
 
"This monster was accepted for preservation in
Kunstkamera and Kosharov was given 4 roubles as a reward"
(there is also, No. 41, February 17, p. 36) (23).
 
(5) Iogani-Georg Dyuvernua (1691-1759) a Russian
 
academician in the department of zoology and anatomy, was
invited to the academy during its foundation. In 1741, by
efforts of Shumakher, he was compelled to leave work and
travel to his homeland, in Germany. Following him, up to
1758 Abraham Kaau-Boerhave (1715-1758) had been in charge
the department of anatomy and physiology in the Academy of
Science. In his era, the anatomical and embryological
collections of the academy came into being, but he practically did not use them. He published one of them in detail
(Abraham Kaau-Boerhave was related to those foreigners,
living in Russia, who did not make anything for promoting
its sciences and for preparing the young Russian scientists (23) .
 
(6) Family of van der Hulst was repeatedly mentioned
in the documents of Petrovskaya epoch. Doctor Zakharii van
der Hulst arrived in Moscow from Holland, apparently in the
70s of the 17th century, and for a long time he was the
physician of Aptekarsky department and court physician at
tsars Ivan and Petr. Then after the death of Ivan, he
became the physician of Petr I. He accompanied him on
both journeys to Arkhangelsk (1693 and 1694). On the
second Journey, he die-' suddenly* . Another van der Hulst
 
 
 
V. Rikhter. "Istoriya meditsiny v Rossii" (History of
Medicine in Russia), p. 2, Moscow, 1820, p. 313,
M. M. Bogoslovskii, Petr I, V. I, p. 181= V. II,
1941, pp. 123-124.
 
 
participated in 1691 in Poteshny battles" in the army
"generalissimusa" of I. I. Buturlin 2 . In 1695 in
"Rospisi nachalnym lyudam Semenovskogo polky" captain
Andrei Yakovlevich van der Hulst was mentioned^. On
the first journey of Petr to Holland, at the Russian
Embassy, lieutenant (or captain) Andrei (Yakovlevich)
van der Hulst 4 became the translator. One year later, t
he was sent by the Dutch government to Moscow as resident*
Later on, the son of the above mentioned "doctor"
Zakharii van der Hulst— Zakhar Zakharovich— was known.
By the order of Peter I, he received "traveller sheet"
via Mozhaisk, Vyazm, Dorogobuzh and Smolensk in foreign
lands for studying science. Although it was impossible
to detect exactly the time of his return to Russia, but
there was no doubt about it . He was a teacher of surgery
at Peterburgsky Hospital 1723 6 . Ya. Chistovich informed,
that "Z. Z. van der Hulst passed the Doctorate degree
examination in Leiden and after returning to Russia he
was the senior doctor in Petersburg Admiral Hospital and,
in addition, a teacher for medical students and pupils of
this hospital. Later on, he lived in Moscow and when a
"Doctor's committee"? (1730) which comprised five doctors
was established, he was one of its members. It is most
 
 
 
2. There also, V. I, p. 127.
 
3. "Sb. vypisok iz arkhivnykh bumag petre Velikom"
(Collection from archives papers about Petr the
 
Great) , VI, Moscow, 1872, p. 148.
 
4. M. M. Bogoslovskii. Petr I, V. II, p. 155, 182, 421.
 
5. There also, V. II, p. 432> V: IV, 1948, p. 252,
339, 340, 344, 346.. M. A. Venevitinov. "Russkie
v Gollandii. Velikoe posol'stvo 1691-1690 godov"
(Russians in Holland. The great embassy 1697-1698)
Moscow, 1897, p. 79.
 
6. V. Rikhter, Istoriya meditsiny v Rossii, p. 3, 1820,
p. 149.
 
7. Ya. Chistovich. "Istoriya pervykh meditsinskikh
shkol v Rossii" (History of first medical schools in Russia) . Appendix X. Alphabetical list of doctors
of medicine, working in Russia in the 18th century.
 
 
probable, that Zakhar Zakharovich van der Hulst is the author
of a dissertation cited in the text on page 7: the difference
of names (Zakharii and Arnold) does not speak against this
supposition, as both names could belong to the same person.
Anyhow there is no doubt about the belonging of the author of
dissertation to the Moscow family of van der Hulst (25) .
 
(7) Speaking about the collection of anatomical preparations, present in Kunstkamera N. G. Kurganov noticed, that
 
"the greatest attention was given to these parts, which explained
parturition. A number of the foeti exceeded more than one
hundred and composed a gradualness from an embryo having the
size of an anisic grain to a completely formed baby. The
collection of monsters was extremely big. These anatomical
descriptions with sketches, reprinted on copper, had a scientific
significance" ("Pismovnik", the second part, p. 196) (32) .
 
(8) In chapter 26, "Gippokratovskoi sbnornik" it is possible to read the following: "All organs are distinguished
simultaneously and they grow, and not one is distinguished
earlier than the other. But the larger ones in nature are
distinguished before the smaller ones, not originating in any
case earlier. However not all receive the final structure
 
in equal time, but some are quicker, others are slower, since
each meets sufficient nutrition. In some, all become distinct
within 40 days in others — within 2 months, in others — within
3 months, and in others — within 4 months" (See V. I. Karnov.
Aristotle i antichnaya embriologiya. Introductory article
in the translation of Aristotle" "0 vozniknovenii zhivotnykh"
Izd. AN SSSR, 1949, p. 23 (35).
 
(9) The embryological opinions of Dekart are stated
 
in the treatise "Opisanie chelovecheskogo telo" (Description
of human body) , where they compose its fourth part — "About
the development of the embryo. Parts, formed in semen" and
the fifth part "the formation of hard parts". The treatise
was published two years before the death of the philosopher,
in 1648. In addition, with the birth conception of living
substances, "which are produced by semen", Dekart also assumed the possibility of spontaneous conception (without
semen and uterus") . *
 
In case of conception from parents, their semen is
mixed and forms cloudy liquid which undergoes a kind of
fermentation, what is formed during this heat widens the
particles of semen, they "press on other particles subsequently locating them gradually. This is the way of
forming the body organs... Heat compels some of the
particles of semen to be collected near definite points
of the space... Thus the heart begins to be formed"2.
The movement of blood from heart makes a way through semen
particles, that is why the blood again returns to the
heart, and by this way the vascular system is formed.
After this, the movement of particles of different kinds
leads to the subsequent formation of the organs - vertebrates with spinal cord, brain, paired organs of sensation
and so on. The energetic character of these embryological
presentations is completely obvious. However, Dekart considered that it is necessary to underline this. "In order
to get acquainted with the figure of the already formed
animal, it should be understood what it represents at the
beginning of its formation and it is necessary to imagine
semen, as some mass, from which, the heart is first formed,
around it the hollow vein is located on one side while on
the other the large artery, united by two tips. The tips
of these vessels, to which the openings of the heart are
directed, indicate the side, where the head must be present,
others also indicate that side, where the lower parts of
the body must be present" 3 (36) .


(10) The idea of preformation may be traced back to
Fr. Balfur, who published more than 70 years ago the first (and for that time excellent) manual book on comparative embryology, in which the Russian works were given proper places (14) .
remote ancient times. Anaksagor taught that "hairs cannot
be formed from no hairs and raft from no raft" similar
ideas were stated by several authors especially Senekoi,  
who in "Questions of nature" wrote: "In semen all future


(2) Samples of these "compositions" are pictured on the prints of that time, reprinted in the book "Kunstkamera Peterbergskoi Akademii Nauk", 1853, 293 p. (see Fig. 6.10 and 11 of the mentioned book) . Description of anatomical and embryological "compositions of Ryuish and texts of the poetical Latin inscriptions which accompanied them much earlier were mentioned by Baer in "Memories about anatomical cabinet" ("Memoire uber das Anatomische Kabinet, gelesen in den Sitzungen der Phys.-math. Kl. d, Sep. 20, 4 u. Oct. 18, 1850; Collection the museum of anthropology and ethnography in Imp. Ac. Sc, 1900, p. 111-152) (21).




1. See S. F. Vasilev. "Evolution ideas in Dekart philosophy (Introductory article in book" Rene Dekart.
Kosmogoniya . Two treatises. GTTI. 1934, p. 121) .


2. R. Dekart. Description of human body. In book:
Here and below figures in round brackets show the number of text page, to which the comment is related.
Kosmogoniya, p. 286-287.  


3. See also pp. 298-299.


(3) Doctor van der Hulst, (see below for data about him) on April 14, 1724 ,; sent organs which belonged to a girl, who died of smallpox, in Kunstkamera. Caesar honored him due to the hermaphrodite (Materials for history of Imp. Ac. Sc, V. 1 Spb, 1855, p. 38) in the other place of these "Materials" wrote due to bringing to the Academy of Science a monster, who was born with two heads, determined by Doctor Stepan Gaokantsk, ...8 roubles were given" (V. II, on March 9, 1732, p. 119) (23).


parts of the human body are contained. The baby in the
(4) In "Materials for history of Imp. Ac. Sc." there are many documents about the obtaining of teratological objects into Kunstkamera.
uterus of the mother has already roots of beard and hair,
which he will carry. In a similar way, in this small mass,
are contained all features of the body as well as all those,
which will be present in his posterity" (see Dzh. Needham,
History of Embryology, page 76) (37) .  


(11) The theory of "investment", conformable to plants
A letter from Vyborkh by colonel general Karnov says "his major Beshentsev Fedor Fedrov in the Vyborgskaya office found a monster of a lamb having four legs and one neck, and the head had the view of a double one. This head had two mouths with tongues and two eyes: they were in the
and animals, was stated by Mal'bransh in the following
expressions: "It seems, although it may be a hardlyaccepted idea, that in one embryo a countless number of trees
is included, because this embryo does not include only a  
tree, serving as its seed, but also, great number of seeds,
which can include new trees and new seeds of trees, containing, in turn, probably in an incomprehensible little form,  
other trees and other seeds, are fruit bearing as the first
and so on till infinity. All what is mentioned about plants
and embryos can be also applied on animals and their embryos,
from which they were produced. In the embryo of a bulb of
tulip, it is possible to distinguish all tulips. Thus in  
the embryo of a fresh non-hatching egg as well, it is possible
to see a chick, which may be nearly completely formed. In
the eggs of frogs, it is easy to recognize frogs, we will
also find other animals in their embryo when we become so
experienced and skilful, that we can open them (N. Mal'bransh.
"seeking out truth" translated by E. B. Smelovaya, V. I, 1903,
p. 51-52) (38) .


(12). Speaking of the influence of the mother's impression on the formation of fetus, Mal'bransh, among other
middle where the forehead must have been " ('Materials..
examples, mentions the following: "One year did not elapse
still from that time, when a woman, looking with great zeal
on the picture of Saint Pia at the time of the celebration of
his canonization, delivered a child, absolutely like the Saint
He had senile face, which was impossible for a child, but did
not possess a beard. His hands were put together as cross
on the chest, his eyes were directed to the sky. He had a
very small forehead, because on the picture, the image of
this Saint was raised to the dome of the church, directed to
the sky, so that his forehead was nearly unnoticed. On his
shoulders, he had something like overturned mitre with some
round birth marks, at these places where mitres were decorated
with precious stones. In short, this infant extremely resembled the picture, but his form was made by the strong
imagination of his mother. All Paris could see him, as I
could, because this infant was preserved for a long time in
spirits of wine" (N. Mal'bransh. Seeking out truth, V. I,
p. 173).  


The description of deformity was so expressive and
V. I, Spb., 1855 No. 165, March 9, 1725, pp. 9-97).
exact, that confidently it was sufficiently possible to
characterize it in terms of recent teratology. Probably,  
the matter was a case of amencephaly or, may be, cerebral
hernia of occipital region. It is not wonderful that the
baby, resembling Saint Pia, was shown in a jar with spirit
directly after bith (38) .  


(13) Philosophy, Leibnits spoke, - gave itself much
Inventory of objects, obtained on March 8, 1725 "Number 1: lamb, with 8 legs, another with three eyes, two trunks, 6 limbs were sent from Tobol'sk by Kozlovsky. Number 2: baby, with 3 legs: from Lower Novgorod from Governor Rzhevsky. Number 3: calf with 2 deformed legs: from Ufa from commandant Bakhmetov. Number 4: baby with two heads also from Bakhmetov. Number 5: one baby, with eyes under the nose and ears under neck: from Nezhin. Number 6 - two babies - breasts and abdomens were joined from Akhtyrok from prince Mikhail Golitsyn; hands, legs and head were normal. Number 7 - baby, with a fish tail, born in Moscow in Tverskaya. Number 8: two puppies, born from a 60-year-old woman, from Akhtyrok from prince Golitsyn. Number 9: baby with 2 heads, 4 hands, 3 legs: from Ufa from commandant Bakhmetov" ("Materials...", V. I, No. 193, March 18, 1625, p. 99).
work about the origin of forms, entelechy and soul. Meanwhile, different accurate investigations, performed on
plants, insects and animals, led to this conclusion: that the  
organic bodies of nature never originated from chaos or not,  
but always from semen, in which, undoubtedly, preformation
was already present... We see something similar, when, for
example, worms become flies and caterpillars become butterflies" (Monadologiya, No. 74), (38).  


(14) "Thus, I suppose that souls which once must
Grigorii Ivanov found a monstrous head puppy with 8 legs ("Materials, V. I, No. 291, September 21, 1725, p. 145).
become human souls, as well as souls of other kinds, existed
in semen, in ancestors up to Adam, i.e. from the very
beginning of things they existed in the form of organic
bodies — a view, which was apparently approved by Swammerdam,
Mal'bransh, Beil', Pitkari, Gartsuker and many other learned
men. This view is also sufficiently confirmed by microscopic
observations of Leeuwenhoek and other fairly prominent
naturalists "(Teoditseya, I, No. 91) (38).  


(15) Leibnits considered that monads are alive and
animated, characterized by incessant change which is accomplished continuously without leaps. "I confirm, as an
indisputable truth — he wrote — that all things were exposed
to change, and became monads and that in each monad this
change was accomplished continuously" (Monadologiya, No. 10) .
From continuous change, the developing monad natural passage
of Leibnits to gradation of monads, forming continuous
eternal ascending series of substances, progressing from unaccomplished to accomplished. By his investigations
"Swammerdam showed that insects, by their respiratory
organs, are similar to plants, and that in nature, an
order of gradualness, descending from animals to plants,
exists. However, there may be, in addition, intermediate
substances between these and others" (Letter of Leibnitz
to Bung). And in another letter: "I am sure, that these
substances must be present, and natural science may discover them. Nature never disturbs continuity anywhere.
It does not make leaps. All categories of substances of
nature form one sole chain, where different classes, like
links, so closely join to each other that for sensual
presentation it is impossible to determine the point, where
each of them begins or ends" (cited by Kuno Fisher)
("History of new philosophy, V. Lebnitz", p. 460) (38) .


(16) About this change of his opinions, Haller wrote
"....Dead body of an infant, with 6 fingers, horn by a daughter of Tikhanov who was working on a marine ship Astrakhan. . .given in Kunstkamera" ("Materials..." V. II, 1886, No. 99, November 29, 1731, p. 83).
the following: "In the body of animal, there is no part
which can originate earlier than the other: all of them
are formed at the same time... If Harvey supposed that


he discovered the epigenetical development, it is because
"On January 17 of this same year, my brother sent a monster which had 2 mouths, the daughter of Ulita Kiryanova the wife of sergeant Nikivor Kosharov gave birth to it: this monster had four hands, four legs and two faces, one had normal view and the other had one eye" ("Materials...", V. Ill, 1886, No. 13, January 20, 1736, p. 14).
from the beginning he only saw small haziness and then the  
rudiments of the head and eyes, exceeding in size all other
bodies, and finally gradually — the internal organs. More
than 20 years ago, i.e. before my numerous observations on
eggs and females of Tetrapoda, I used this argument to prove
that embryo strongly differed from the formed animal, while
I confirmed, that in animals at the moment of conception
parts which were present in completely formed animal were
absent. From this time, I had the complete possibility to
confirm that all that was deduced by me against preformation
theory, in fact speaks in its favor" (cited by Dzh. Needham,  
History of Embryology, p. 226-227) (39).  


(17) On the basis of the confirmation of the Bible,
"This monster was accepted for preservation in Kunstkamera and Kosharov was given 4 roubles as a reward" (there is also, No. 41, February 17, p. 36) (23).
that earth and mankind populating it, have existed about
6000 years and from that the average duration of life of


man is equal to 30 years, Haller calculated, that God created
(5) Iogani-Georg Dyuvernua (1691-1759) a Russian academician in the department of zoology and anatomy, was invited to the academy during its foundation. In 1741, by efforts of Shumakher, he was compelled to leave work and travel to his homeland, in Germany. Following him, up to 1758 Abraham Kaau-Boerhave (1715-1758) had been in charge the department of anatomy and physiology in the Academy of Science. In his era, the anatomical and embryological collections of the academy came into being, but he practically did not use them. He published one of them in detail (Abraham Kaau-Boerhave was related to those foreigners, living in Russia, who did not make anything for promoting its sciences and for preparing the young Russian scientists (23) .
at the same time a minimum of 2000,000,000,000 people (cited
by article of Kirchhoff on Wolff, p. 204) (39) .  


(18) Speaking about the impression produced on contemporaries by the discovery of Bfennet (about the development of bodies from unfertilized ovum), A. E. Gaisinovich noticed
(6) Family of van der Hulst was repeatedly mentioned in the documents of Petrovskaya epoch. Doctor Zakharii van der Hulst arrived in Moscow from Holland, apparently in the 70s of the 17th century, and for a long time he was the physician of Aptekarsky department and court physician at tsars Ivan and Petr. Then after the death of Ivan, he became the physician of Petr I. He accompanied him on both journeys to Arkhangelsk (1693 and 1694). On the second Journey, he die-' suddenly* . Another van der Hulst participated in 1691 in Poteshny battles" in the army "generalissimusa" of I. I. Buturlin 2 . In 1695 in "Rospisi nachalnym lyudam Semenovskogo polky" captain Andrei Yakovlevich van der Hulst was mentioned^. On the first journey of Petr to Holland, at the Russian Embassy, lieutenant (or captain) Andrei (Yakovlevich) van der Hulst 4 became the translator. One year later, t he was sent by the Dutch government to Moscow as resident* Later on, the son of the above mentioned "doctor" Zakharii van der Hulst— Zakhar Zakharovich— was known. By the order of Peter I, he received "traveller sheet" via Mozhaisk, Vyazm, Dorogobuzh and Smolensk in foreign lands for studying science. Although it was impossible to detect exactly the time of his return to Russia, but there was no doubt about it . He was a teacher of surgery at Peterburgsky Hospital 1723 6 . Ya. Chistovich informed, that "Z. Z. van der Hulst passed the Doctorate degree examination in Leiden and after returning to Russia he was the senior doctor in Petersburg Admiral Hospital and, in addition, a teacher for medical students and pupils of this hospital. Later on, he lived in Moscow and when a "Doctor's committee"? (1730) which comprised five doctors was established, he was one of its members. It is most probable, that Zakhar Zakharovich van der Hulst is the author of a dissertation cited in the text on page 7: the difference of names (Zakharii and Arnold) does not speak against this supposition, as both names could belong to the same person. Anyhow there is no doubt about the belonging of the author of dissertation to the Moscow family of van der Hulst (25) .
that the scientists of that time... lost sight of that it
proved in the best case only ovism but not preformation
(Cited article in the edition of translation "Theory of  
conception" of Wolff, 1950, p. 379). With the latter confirmation, it is difficult to agree: ovism is one of two
forms of the preformation theory and is principally
identical with its other form — animalculism. (40)  


(19) After that the chapters of this book, dedicated
by Wolff, were written, he mentioned in the preface
"Theory of conception" of Wolff (Publisher House Ac. Sc.
USSR, 1950) which was published with the supplement of
article A. E. Gaisinovich "K. F. Wolff and studies on
development" (pp. 363-477) Tasks, which the author of this
interesting article put before himself, did not allow him,
apparently, to stop and dwell in more detail on embryological and teratological works of Wolff.


Here it is also necessary to notice that the translation replaced in the Russian edition of Wolff the term
V. Rikhter. "Istoriya meditsiny v Rossii" (History of Medicine in Russia), p. 2, Moscow, 1820, p. 313, M. M. Bogoslovskii, Petr I, V. I, p. 181= V. II, 1941, pp. 123-124.
"generation" by the word "conception", which could not be
considered felicitous. The term "conception" meant the
beginning stage of development, appearance of a new individual, while Wolff did not mean only this stage, but also all
subsequent individual development. It is completely accurate
to translate "generation" by the word "development". Wolff
himself used in his German book neither the term "conception"
(Entstehung) nor the term "development" (Entwickelung) , and
dept the Latin root in the German word "die Generation". In
accordance with this, upon the examination of Wolff's dissertation and its popular summary in German below the original
Wolff term "generation" is preserved (43) .  


(20) In the first volume of "Zur Morphologie" Gete
2. There also, V. I, p. 127.
mentioned Wolff in four places. He gave a brief account of
biography of Wolff (p. 80-83), mentioned in the notes of
Murzinitsa on Wolff (p. 252-256), then dwelt on studies of
Wolff about metamorphosis of plants (p. 83-87) . At last he
tackled the understanding of educational yearning, using the
terms of Blumenbakh (pp. 114-116).  


In the last extract Gete writes the following "In
3. "Sb. vypisok iz arkhivnykh bumag petre Velikom" (Collection from archives papers about Petr the Great) , VI, Moscow, 1872, p. 148.
criticism of ability to reach an opinion" Kant states "In relation to the theory of epigensis no one worked either
for its proof and for substantiation of true principles of
its application, or partly for the restriction of its
extremely wide application, as Mr. Buildings advisor
Blumenbakh". This evidence of honest Kant encouraged me
again to scrutinize work of Blumenbakh, which I truly read
before, but did not imbue it. Here I found my Christof
(1-L.B.) Friedrich Wolff as an intermediate link between
Haller and Bonnet; on the one hand, and Blumenbakh, on the
other. For his epigenesis, Wolff must suppose the presence
of an organic element, on which creatures feed, which is
intended for organic life, and supplied this material by an
essential force". The mentioned words of Gete witness to
the superficial acquaintance of the great poet with the
views of Wolff (Even Wolff's name was written incorrectly
by Gete (44) .  


(21) A. E. Gaisinovich (1950, p. 462-463) mentioned
4. M. M. Bogoslovskii. Petr I, V. II, p. 155, 182, 421.
information, that Wolff at the beginning of the 70th year
taught in the academic high school— chemistry, anatomy and
botany, and he also directed the preparation of the scientific activity of student Fedor Galchenkov (48) .  


(22) This place in the translation of Meckel tendentiously stated: instead of "the Highest Creator" was put
5. There also, V. II, p. 432> V: IV, 1948, p. 252, 339, 340, 344, 346.. M. A. Venevitinov. "Russkie v Gollandii. Velikoe posol'stvo 1691-1690 godov" (Russians in Holland. The great embassy 1697-1698) Moscow, 1897, p. 79.
"The creating nature". Wolff disputed the preformation of
formulated parts; in the German translation, his skepticism
is not related to the Creator for the authority is untouched,  
but to nature. This "liberty of translation" bars the radicality of scientific and philosophical view of Wolff (76) .  


(23) Here the phrase made by Wolff, has no connection
6. V. Rikhter, Istoriya meditsiny v Rossii, p. 3, 1820, p. 149.
with other discussions and clearly intended for not blaming
atheism: eiusmodi vero materia, talibus, viribus instructa
immediate a Deo ex nihilo creata sit (it is also true that
material, supplied by these forces, is directly created by
God from nothingness) (76) .  


(24) A. E. Gaisinovich (1950, p. 455) repeated the
7. Ya. Chistovich. "Istoriya pervykh meditsinskikh shkol v Rossii" (History of first medical schools in Russia) . Appendix X. Alphabetical list of doctors of medicine, working in Russia in the 18th century.
mentioned statement of K. M. Baer nearly literally: "This
remarkable work of Wolff... did not draw the attention of  
all the scientific world up to 1812, when Meckel translated
it from the Latin language". It is necessary to notice,
however, that the work of Wolff was given due attention and
it was evaluated as a remarkable work in the book of
I. Bezeke, published in 1797 (see p. 114) and in the dissertation L. Tredern (1808) (see Chapter 11) (87) .  


(25) Apparently, in the declaration of biological
works by the Academy of Science for a prize, Wolff showed
initiatives also early. Thus, in 1779 the academy declared
a competition for a prize on the question about reproduction
of cryptogamous plants. This theme of competitive work


was written in expressions which a great probability impel
(7) Speaking about the collection of anatomical preparations, present in Kunstkamera N. G. Kurganov noticed, that
to consider Wolff the author of this question: "Theoriam
generationis et fructificationis plantarum cryptogamicarum
Linnaei dare etc...." (gives the theory of development and
fruiting of cryptogamous plants, by system of Linnae, plants
and so on) .


In 1783, this prize was awarded to a professor in
"the greatest attention was given to these parts, which explained parturition. A number of the foeti exceeded more than one hundred and composed a gradualness from an embryo having the size of an anisic grain to a completely formed baby. The collection of monsters was extremely big. These anatomical descriptions with sketches, reprinted on copper, had a scientific significance" ("Pismovnik", the second part, p. 196) (32) .
Leipzig, log. Gedvig for the work under the title "Theoria
generationis et fructificationis plantarum cryptogamicarum
Lennaei, mere propriis observationibus et experimentis
superstructa dessertatio, quae praemio ab. Academia imp.
Petropol. pro Anno MDCCLXXXIII proposito ornata est" (105).  


(26) It is necessary to notice, that the embryological
(8) In chapter 26, "Gippokratovskoi sbnornik" it is possible to read the following: "All organs are distinguished simultaneously and they grow, and not one is distinguished earlier than the other. But the larger ones in nature are distinguished before the smaller ones, not originating in any case earlier. However not all receive the final structure in equal time, but some are quicker, others are slower, since each meets sufficient nutrition. In some, all become distinct within 40 days in others — within 2 months, in others " within 3 months, and in others " within 4 months" (See V. I. Karnov. Aristotle i antichnaya embriologiya. Introductory article in the translation of Aristotle" "0 vozniknovenii zhivotnykh" Izd. AN SSSR, 1949, p. 23 (35).
problems have interested Petersburg Academy of Science before  
the arrival of Wolff. It is possible to judge this, in  
particular, by the published collection in 1756, which is
composed of two works (dissertations) , and sent to the
competition announced by the academy. The question, put by
the academy, touched upon the possible influence of the
impression, felt by the pregnant woman, on the developing
fetus. One of the mentioned works belonged to a professor


at Leipzig University C. Ch. Krauze*, and another — to a
(9) The embryological opinions of Dekart are stated in the treatise "Opisanie chelovecheskogo telo" (Description of human body) , where they compose its fourth part — "About the development of the embryo. Parts, formed in semen" and the fifth part "the formation of hard parts". The treatise was published two years before the death of the philosopher, in 1648. In addition, with the birth conception of living substances, "which are produced by semen", Dekart also assumed the possibility of spontaneous conception (without semen and uterus") . *
member of Petersburg Academy of Science, I. G. Rederer.  
In lively controversy on this theme Byuffon participated.
He energetically objected to preformists, who according to
their views, the embryo is proved to be similar with the  
parents under the influence of imagination of the mother.  


In case of conception from parents, their semen is mixed and forms cloudy liquid which undergoes a kind of fermentation, what is formed during this heat widens the particles of semen, they "press on other particles subsequently locating them gradually. This is the way of forming the body organs... Heat compels some of the particles of semen to be collected near definite points of the space... Thus the heart begins to be formed"2. The movement of blood from heart makes a way through semen particles, that is why the blood again returns to the heart, and by this way the vascular system is formed. After this, the movement of particles of different kinds leads to the subsequent formation of the organs - vertebrates with spinal cord, brain, paired organs of sensation and so on. The energetic character of these embryological presentations is completely obvious. However, Dekart considered that it is necessary to underline this. "In order to get acquainted with the figure of the already formed animal, it should be understood what it represents at the beginning of its formation and it is necessary to imagine semen, as some mass, from which, the heart is first formed, around it the hollow vein is located on one side while on the other the large artery, united by two tips. The tips of these vessels, to which the openings of the heart are directed, indicate the side, where the head must be present, others also indicate that side, where the lower parts of the body must be present" 3 (36) .


(10) The idea of preformation may be traced back to remote ancient times. Anaksagor taught that "hairs cannot be formed from no hairs and raft from no raft" similar ideas were stated by several authors especially Senekoi, who in "Questions of nature" wrote: "In semen all future parts of the human body are contained. The baby in the uterus of the mother has already roots of beard and hair, which he will carry. In a similar way, in this small mass, are contained all features of the body as well as all those, which will be present in his posterity" (see Dzh. Needham, History of Embryology, page 76) (37) .


Caroli Christiani Krauze. Dissertatio de questione ab
1. See S. F. Vasilev. "Evolution ideas in Dekart philosophy (Introductory article in book" Rene Dekart. Kosmogoniya . Two treatises. GTTI. 1934, p. 121) .
Academia imp. scientianim petropolitana pro praemis in  
annum MOCCLVI proposita. Quaenam sit causa proxima
mutans, corpus foetus etc... There also, J. D. Roederer
Dessertatis Petropoli 1956.  


2. R. Dekart. Description of human body. In book: Kosmogoniya, p. 286-287.


In the controversy Terner also participated. He objected to
3. See also pp. 298-299.
Blondel, denying the influence of mother imagination on the
embryos. Argument of Terner was confirmed by the fact that
the blood vessels of the mother directly passes in the vessels
of the fetus. Ens subscribed to Terner' s opinion. Apparently,
the difference, existing on this question in literature,
induced Petersburg Academy of Science to announce competition
on works which would comprehensively answer it. The scientific
committee of the Academy, examining the competitive works,
revealed scientific impartiality, and the Academy published
two works, each one of them represented a contradictory opinion.  
Krauze was a supporter, and Rederer was an opponent of the
influence of maternal imagination on the fetus.  


Krauze considered that the question raised was very
difficult, nearly hopeless to solve. However, he noticed,
that many examples were present, when the fetus was changed
in a way, that not only simple people and the mother herself,
but even sharply sensitive scientific people in medical practice
related it, although partially, to the strong emotional shock
of the mother. Enumerating the opinions of authors, discussing
this question (Sennert, Morisso, Ludvig, Hofman, Abraham
Kau-burgav, Takhoni and others), Krauze suggested, that the
reader can make for himself the most classical ironical
defiance: "Hie Rhodus, hie salta!" (literally: "Here
Rodos also jump!") and all the following statements attempt
to answer this call. As also the majority of his predecessors, Krauze paid attention mainly to cases of appearance of
pigmental birth marks, where their forms and situation compel
to suggest the influence of pregnant women. He began with
these sharp effects, as terror, fear, anger and so on, showing strong physiological influence expressed first of all in
the reaction from the side of vascular and nervous systems.
From this Krauze made the conclusion, that "if the brain was
strongly alarmed, then small changes took place in the body".
Referring to known cases of adult people suddently growing
grey under the effect of deep feelings, Krauze suggested that,
there were more bases to expect changes of skin color of the
fetus under the effect of maternal imagination. If terror
or fear can cause small ulcers on lips or erysipelas, then
why does it seem incredible to you, that the same phenomena
can take place in the body of the fetus, whose structure is
so weak and delicate, and its vessels are so numerous and
full of juices?"


611
(11) The theory of "investment", conformable to plants and animals, was stated by Mal'bransh in the following expressions: "It seems, although it may be a hardlyaccepted idea, that in one embryo a countless number of trees is included, because this embryo does not include only a tree, serving as its seed, but also, great number of seeds, which can include new trees and new seeds of trees, containing, in turn, probably in an incomprehensible little form, other trees and other seeds, are fruit bearing as the first and so on till infinity. All what is mentioned about plants and embryos can be also applied on animals and their embryos, from which they were produced. In the embryo of a bulb of tulip, it is possible to distinguish all tulips. Thus in the embryo of a fresh non-hatching egg as well, it is possible to see a chick, which may be nearly completely formed. In the eggs of frogs, it is easy to recognize frogs, we will also find other animals in their embryo when we become so experienced and skilful, that we can open them (N. Mal'bransh. "seeking out truth" translated by E. B. Smelovaya, V. I, 1903, p. 51-52) (38) .


(12). Speaking of the influence of the mother's impression on the formation of fetus, Mal'bransh, among other examples, mentions the following: "One year did not elapse still from that time, when a woman, looking with great zeal on the picture of Saint Pia at the time of the celebration of his canonization, delivered a child, absolutely like the Saint He had senile face, which was impossible for a child, but did not possess a beard. His hands were put together as cross on the chest, his eyes were directed to the sky. He had a very small forehead, because on the picture, the image of this Saint was raised to the dome of the church, directed to the sky, so that his forehead was nearly unnoticed. On his shoulders, he had something like overturned mitre with some round birth marks, at these places where mitres were decorated with precious stones. In short, this infant extremely resembled the picture, but his form was made by the strong imagination of his mother. All Paris could see him, as I could, because this infant was preserved for a long time in spirits of wine" (N. Mal'bransh. Seeking out truth, V. I, p. 173).


The description of deformity was so expressive and exact, that confidently it was sufficiently possible to characterize it in terms of recent teratology. Probably, the matter was a case of amencephaly or, may be, cerebral hernia of occipital region. It is not wonderful that the baby, resembling Saint Pia, was shown in a jar with spirit directly after bith (38) .


In order to imagine the possibility of this phenomenon,  
(13) Philosophy, Leibnits spoke, - gave itself much work about the origin of forms, entelechy and soul. Meanwhile, different accurate investigations, performed on plants, insects and animals, led to this conclusion: that the organic bodies of nature never originated from chaos or not, but always from semen, in which, undoubtedly, preformation was already present... We see something similar, when, for example, worms become flies and caterpillars become butterflies" (Monadologiya, No. 74), (38).
which is shown by the influence of changes in the organism
of the mother on the fetus, it is necessary to prove their
close relation. On his side Krauze confirmed, "fetus with
the uterus represents a single continuous whole". According to this opinion, this is especially related to the
nervous system, so that "stimulation of the nerves of the
uterus may and must be passed to all the nervous system
of the fetus. Distributing this confirmation on psychics,
as well as suggesting, that the fetus was capable of  
psycological manifestations, Krauze deduced the following
conclusion: "In the brain of the fetus exists the same
condition, which exists in the brain of the mother". All
these views are summarized in the concluding paragraph of
the dissertation". What was stated so far may be added to
all these examples, which instruct, that the fetus body is
changed from the mother, if her soul is strongly shocked.  
Ideas, originating in the mother's brain are "united with
fetus brain"; they are stimulated in it more quickly and
energetically, than in the soul of the mother herself, as
the pulse of arteries and effect of nervous and generally
all responses in babies are quicker than in adults. Under
the influence of these ideas, the fetus brain gives effect
on its body and namely in this way, which corresponds to
perceived ideas. Therefore, the fetus's brain produces in
the corresponding parts of his body the same things which
he himself undergoes".  


Directly after the considered work of Krauze, it is
(14) "Thus, I suppose that souls which once must become human souls, as well as souls of other kinds, existed in semen, in ancestors up to Adam, i.e. from the very beginning of things they existed in the form of organic bodies — a view, which was apparently approved by Swammerdam, Mal'bransh, Beil', Pitkari, Gartsuker and many other learned men. This view is also sufficiently confirmed by microscopic observations of Leeuwenhoek and other fairly prominent naturalists "(Teoditseya, I, No. 91) (38).
necessary to examine the work of Rederer, which is simply
entitled "Dissertatis" and also begins with the formulation
of the question raised by the Academy: "What is the
direct cause for changing the fetus body?..."


In order to possibly answer this stated question,
(15) Leibnits considered that monads are alive and animated, characterized by incessant change which is accomplished continuously without leaps. "I confirm, as an indisputable truth — he wrote — that all things were exposed to change, and became monads and that in each monad this change was accomplished continuously" (Monadologiya, No. 10) . From continuous change, the developing monad natural passage of Leibnits to gradation of monads, forming continuous eternal ascending series of substances, progressing from unaccomplished to accomplished. By his investigations "Swammerdam showed that insects, by their respiratory organs, are similar to plants, and that in nature, an order of gradualness, descending from animals to plants, exists. However, there may be, in addition, intermediate substances between these and others" (Letter of Leibnitz to Bung). And in another letter: "I am sure, that these substances must be present, and natural science may discover them. Nature never disturbs continuity anywhere. It does not make leaps. All categories of substances of nature form one sole chain, where different classes, like links, so closely join to each other that for sensual presentation it is impossible to determine the point, where each of them begins or ends" (cited by Kuno Fisher) ("History of new philosophy, V. Lebnitz", p. 460) (38) .
Rederer considered that it is necessary to study thoroughly,  
how the mother's body unites with the embryo, taking into
consideration, that the only connection between them is
carried out through the placenta. The investigation of
this connection led him to the following conclusion. The
lumens of the blood vessels of the placenta, directed to
the uterus, up till this stage, are so narrow. They do
not admit turpentine oil or any other liquid to pass.  
True anastomoses, connecting the vessels of the uterus


612
(16) About this change of his opinions, Haller wrote the following: "In the body of animal, there is no part which can originate earlier than the other: all of them are formed at the same time... If Harvey supposed that


he discovered the epigenetical development, it is because from the beginning he only saw small haziness and then the rudiments of the head and eyes, exceeding in size all other bodies, and finally gradually — the internal organs. More than 20 years ago, i.e. before my numerous observations on eggs and females of Tetrapoda, I used this argument to prove that embryo strongly differed from the formed animal, while I confirmed, that in animals at the moment of conception parts which were present in completely formed animal were absent. From this time, I had the complete possibility to confirm that all that was deduced by me against preformation theory, in fact speaks in its favor" (cited by Dzh. Needham, History of Embryology, p. 226-227) (39).


(17) On the basis of the confirmation of the Bible, that earth and mankind populating it, have existed about 6000 years and from that the average duration of life of


and vessels of placenta, do not exist; Vasa hypogastrica
man is equal to 30 years, Haller calculated, that God created at the same time a minimum of 2000,000,000,000 people (cited by article of Kirchhoff on Wolff, p. 204) (39) .
which is full, by all means of a waxen mass or any other
fluid, does not pass its contents to the umbilical vessels.
This was established either in human carcass, or in living
cows, ewes, bitches and other animals. If the pregnant
animal is fed on roots of Rabiae tinctorum, then the fetus
bones will not be stained with the red color. The blood
of the fetus differs from the mother's blood in that it is
more liquid. Finally, the fetus pulse also differs from
the mother's rate.  


Subsequently, the fetus lives its independent life,
(18) Speaking about the impression produced on contemporaries by the discovery of Bfennet (about the development of bodies from unfertilized ovum), A. E. Gaisinovich noticed that the scientists of that time... lost sight of that it proved in the best case only ovism but not preformation (Cited article in the edition of translation "Theory of conception" of Wolff, 1950, p. 379). With the latter confirmation, it is difficult to agree: ovism is one of two forms of the preformation theory and is principally identical with its other form — animalculism. (40)
moving its own blood by its forces, beating of its heart,
by blood circulation and through placenta, without the  
help of the maternal blood. Therefore, rest, movement,
sleeping, awakening and even life and death of the mother
and fetus are not obligatory present in harmony. Later on,  
Rederer categorically denied the existence of a nervous
connection between the placenta and the mother. The
comparison of the positions of the birth marks as well as
their presence or absence in the mother and baby does not
give, according to his opinion, basis to suggest the
influence through blood or nervous system. Rederer discussed in detail the question about "mind" of the fetus
and passed to the conclusion, that this mind cannot
adequately respond to the feeling of the mother. This is
stated by the examples of strong shocks in the mother, not  
accompanied by the appearance of birth marks in the baby,
and examples of appearance of birth marks and warts without
any connection with the mother's feeling. In the following
pages the various deformities of human beings are mentioned,  
and Rederer reached the conclusion that the birth marks
differ from true deformities only quantitatively. They are
also disturbances of the normal development. As the
deformities frequently affect the internal organs, then in
relation to them, the usual supposition about maternal
influence loses its significance. The general conclusion
of Rederer comes to that the confirmations about the
influence of the mother's feeling on the baby are not
supported by verified facts, and are but the product of
fantasy (105) .


(27) The effect of vitalistic views of Blumenbackh on
(19) After that the chapters of this book, dedicated by Wolff, were written, he mentioned in the preface "Theory of conception" of Wolff (Publisher House Ac. Sc. USSR, 1950) which was published with the supplement of article A. E. Gaisinovich "K. F. Wolff and studies on development" (pp. 363-477) Tasks, which the author of this interesting article put before himself, did not allow him, apparently, to stop and dwell in more detail on embryological and teratological works of Wolff.
some Russian physicians-biologists can be traced up till the  
20 's of the 19th century. Thus, in 1825 at Moscow University, the surgeon Nikifor Dmitrievich Lebedev discussed a thesis for the degree of Doctor of Medicine. He
later on read at the university "history and literature of  
medicine". Lebedev dissertation was entitled "About the
nature of weightless substances in general and vital
powers in particular" (Dissertatio inauguralis physislogica
de natura imponderab ilium in genere et de viribus vitalibus
in specie, quam...in Universitate caesarea Mosquensi, pro
gradu doctoris medicinae. . .elaboravit publiceque defendet
chirurgus I-mae classis Nicephorus Lebedev, Mosquae, 1825.  
28 p.). Lebedev, referring to Blumenbakh, dwelt on the
idea that all vital processes — organic formation and growth
as well as movement of already formed parts-possess their
own source of a special vital power (55) . The vital power,
according to his opinion, is an internal, inherent character
in the organic body, which is the cause of life and at the
same time seems to be its product (Vis vitalis est interna
et proporia organici corpori qualitis, quae vitae causam
constituit et simul ejusdem est quasi productum, thesis 9) .
Works of similar kind are not characteristic of the common
materialistic trend of Russian biological and medical
sciences. Therefore, the dissertation of Lebedev served
as a reference and mainly as an illustration of this
negative influence, which was shown by the idealistic
German philosophy and some representatives of Russian
sciences especially the naturalists who worked under its
direct influence (106) .  


(28) Izef Gotlib Kelreiter (1733-1806), botanist and
Here it is also necessary to notice that the translation replaced in the Russian edition of Wolff the term "generation" by the word "conception", which could not be considered felicitous. The term "conception" meant the beginning stage of development, appearance of a new individual, while Wolff did not mean only this stage, but also all subsequent individual development. It is completely accurate to translate "generation" by the word "development". Wolff himself used in his German book neither the term "conception" (Entstehung) nor the term "development" (Entwickelung) , and dept the Latin root in the German word "die Generation". In accordance with this, upon the examination of Wolff's dissertation and its popular summary in German below the original Wolff term "generation" is preserved (43) .
zoologist, worked from 1756 to 1761 as a junior scientific
assistant at Petersburg Academy of Sciences, with which he
kept in close contact after returning back to Germany and
till the end of his life. In the period, from 1758 to 1811,  
15 botanical and more than 20 zoological works of Kelreiter
were published in Russian scientific and scientific popular
editions (Novi Commentarii Acad. Scient. petropolitanae,  
Acta Acad. Scient. petropol. Nova Acta Ac. Sc. Petropl.,  
Trudy Volnogo Ekonomicheskogo Obshchestva and in the journal
"Sochineniya,K poize i uveceleniyu sluzhashchie") . His
scientific fame is connected mainly with the study of  
reproduction and hybridization in plants (see Ioz. Kelreiter"  
Study about sex and hybridization in the plant", the editor
with a biological essay was prof. E. V. Wolff, 1840).  


(20) In the first volume of "Zur Morphologie" Gete mentioned Wolff in four places. He gave a brief account of biography of Wolff (p. 80-83), mentioned in the notes of Murzinitsa on Wolff (p. 252-256), then dwelt on studies of Wolff about metamorphosis of plants (p. 83-87) . At last he tackled the understanding of educational yearning, using the terms of Blumenbakh (pp. 114-116).


In the last extract Gete writes the following "In criticism of ability to reach an opinion" Kant states "In relation to the theory of epigensis no one worked either for its proof and for substantiation of true principles of its application, or partly for the restriction of its extremely wide application, as Mr. Buildings advisor Blumenbakh". This evidence of honest Kant encouraged me again to scrutinize work of Blumenbakh, which I truly read before, but did not imbue it. Here I found my Christof (1-L.B.) Friedrich Wolff as an intermediate link between Haller and Bonnet; on the one hand, and Blumenbakh, on the other. For his epigenesis, Wolff must suppose the presence of an organic element, on which creatures feed, which is intended for organic life, and supplied this material by an essential force". The mentioned words of Gete witness to the superficial acquaintance of the great poet with the views of Wolff (Even Wolff's name was written incorrectly by Gete (44) .


614
(21) A. E. Gaisinovich (1950, p. 462-463) mentioned information, that Wolff at the beginning of the 70th year taught in the academic high school— chemistry, anatomy and botany, and he also directed the preparation of the scientific activity of student Fedor Galchenkov (48) .


(22) This place in the translation of Meckel tendentiously stated: instead of "the Highest Creator" was put "The creating nature". Wolff disputed the preformation of formulated parts; in the German translation, his skepticism is not related to the Creator for the authority is untouched, but to nature. This "liberty of translation" bars the radicality of scientific and philosophical view of Wolff (76) .


(23) Here the phrase made by Wolff, has no connection with other discussions and clearly intended for not blaming atheism: eiusmodi vero materia, talibus, viribus instructa immediate a Deo ex nihilo creata sit (it is also true that material, supplied by these forces, is directly created by God from nothingness) (76) .


The work of Kelreiter about the irritability in the  
(24) A. E. Gaisinovich (1950, p. 455) repeated the mentioned statement of K. M. Baer nearly literally: "This remarkable work of Wolff... did not draw the attention of all the scientific world up to 1812, when Meckel translated it from the Latin language". It is necessary to notice, however, that the work of Wolff was given due attention and it was evaluated as a remarkable work in the book of I. Bezeke, published in 1797 (see p. 114) and in the dissertation L. Tredern (1808) (see Chapter 11) (87) .
plant, to which Wolff referred, was called: "Nouvelles
observations et experiences sur l'irritabilite des etamines
de l'epine vinette (Berberis vulgaris)", Traduit de l'allemand
par M. I 1 adjoint Sewergin, Nova Acta Acad. Scient. Petropol.,
4, 1790 (German original was received in Petersburg in
1788) (112).  


(29) The point of view of A. E. Gaisinovich is
(25) Apparently, in the declaration of biological works by the Academy of Science for a prize, Wolff showed initiatives also early. Thus, in 1779 the academy declared a competition for a prize on the question about reproduction of cryptogamous plants. This theme of competitive work
similar to the presentation mentioned here about the outlook of Wolff (see Wolff "Theory of conception") . It is
possible to combine with it the confirmation, that the  
studies on preformation and epigenesis do not always correspond to the demarcation between idealism and materialism
in biology. The following serves as an evidence, that
preformists were also idealists (Leibnitz, Haller, Bonnet)
and mechanical materialists (for example, Lamettri) ; equally
as epigenetics, were either idealists (Aristotle Harvei) ,


or mechanical materialists (Dekart, Maupertuis, Byuffon and
was written in expressions which a great probability impel to consider Wolff the author of this question: "Theoriam generationis et fructificationis plantarum cryptogamicarum Linnaei dare etc...." (gives the theory of development and fruiting of cryptogamous plants, by system of Linnae, plants and so on) .
Didar3t).A. E. Gaisinovich included Wolff as well in the
latter category. His "absolute epigenesis" was grouped
together with Gaisinovich in the mechanical materialism.
For its complete verification, this statement could be
desired as only a less categorical expression. In the
evaluation of Wolff's outlook, it is impossible, apparently,
to deny absolutely his fluctuations between materialism and  
idealism; these fluctuations were an unavoidable originality
even by the most prominent thinkers of all historical epochs,  
preceding the formation of successive system, i.e. dialectic
system and materialism (119) .  


(30) Semen Gerasimovich Zybelin (1735-1802), after
In 1783, this prize was awarded to a professor in Leipzig, log. Gedvig for the work under the title "Theoria generationis et fructificationis plantarum cryptogamicarum Lennaei, mere propriis observationibus et experimentis superstructa dessertatio, quae praemio ab. Academia imp. Petropol. pro Anno MDCCLXXXIII proposito ornata est" (105).
finishing the study in Moscow Ecclesiastical Academy, joined
Moscow University till the opening there of a medical
faculty. After he finished the course, he was sent to study
medicine abroad. At Leiden, Zybelin defended a doctor's
dissertation in 1764. After returning back to Moscow he
read courses of anatomy, physiology, chemistry, pharmacology
and therapy as a professor in the mediqal faculty. In 1784,
he was elected member of the Academy of Science (121) .  


(31) Prince Dmitrii Alekseevich Golitsyn (1731-1803),  
(26) It is necessary to notice, that the embryological problems have interested Petersburg Academy of Science before the arrival of Wolff. It is possible to judge this, in particular, by the published collection in 1756, which is composed of two works (dissertations) , and sent to the competition announced by the academy. The question, put by the academy, touched upon the possible influence of the impression, felt by the pregnant woman, on the developing fetus. One of the mentioned works belonged to a professor
a prominent Russian diplomat, was a former ambassador in Holland and France, friend of Diderot and Galvetsy, known as
author of many physical works (122) .  


(32) Dzh. Needham in his "History of Embryology"
at Leipzig University C. Ch. Krauze*, and another — to a member of Petersburg Academy of Science, I. G. Rederer. In lively controversy on this theme Byuffon participated. He energetically objected to preformists, who according to their views, the embryo is proved to be similar with the parents under the influence of imagination of the mother.
prefaced the list of the literature sources used by him with
a list of works, which he could not obtain. As "less
important works on the history of embryology" Needham also
mentioned Bezeke's book. It is difficult to decide, on
which basis the English historian of embryology considered
Bezeke's book less important, if we take into consideration
that according to his confession, he never saw it (122) .  


(33) Two extracts compose the contents of the already
Caroli Christiani Krauze. Dissertatio de questione ab Academia imp. scientianim petropolitana pro praemis in annum MOCCLVI proposita. Quaenam sit causa proxima mutans, corpus foetus etc... There also, J. D. Roederer Dessertatis Petropoli 1956.
mentioned book of 1797 (in it, as stated, were present an
essay on the history of a hypothesis about the conception
and development of animals and in addition, "History of the
origin of division of the natural bodies into three kingdoms")
The third extract appeared in the form of a separate small
volume in the year of the author's death (J. M. G. Bezeke.  
Allgemeine Geschichte der Naturgeschichte in dem Zeitraume
von Erschaffung der Welt bis auf das Jahr N. C. G. 1791.  
Mitau, 1802, XXXII±154S) .  


For more details about the works of Bezeke see the
article by the author of the present book in "Trudy instituta
istoriii estestvoznaniya i tekhniki, V.IV, 1955" (Works of
the Institute of History of Natural Sciences and Technique)
(123).


(34) Matvei Khristianovich Peken was born in Petersburg,
In the controversy Terner also participated. He objected to Blondel, denying the influence of mother imagination on the embryos. Argument of Terner was confirmed by the fact that the blood vessels of the mother directly passes in the vessels of the fetus. Ens subscribed to Terner' s opinion. Apparently, the difference, existing on this question in literature, induced Petersburg Academy of Science to announce competition on works which would comprehensively answer it. The scientific committee of the Academy, examining the competitive works, revealed scientific impartiality, and the Academy published two works, each one of them represented a contradictory opinion. Krauze was a supporter, and Rederer was an opponent of the influence of maternal imagination on the fetus.
he studied medicine in Ien, where he obtained the degree of  
Doctor of Medicine (according to other data-- in Gettingen) .  
When he returned back to Russia he worked as the admiralty
doctor, read a course of obstetrics in Petersburg hospitals.  
From 1793, he travelled to Moscow, where he read pathology
and organized the first therapeutic clinic with ten beds (124)


(35) Nestor Maksimovich Maksimovich-Ambodic (1744-1812)  
Krauze considered that the question raised was very difficult, nearly hopeless to solve. However, he noticed, that many examples were present, when the fetus was changed in a way, that not only simple people and the mother herself, but even sharply sensitive scientific people in medical practice related it, although partially, to the strong emotional shock of the mother. Enumerating the opinions of authors, discussing this question (Sennert, Morisso, Ludvig, Hofman, Abraham Kau-burgav, Takhoni and others), Krauze suggested, that the reader can make for himself the most classical ironical defiance: "Hie Rhodus, hie salta!" (literally: "Here Rodos also jump!") and all the following statements attempt to answer this call. As also the majority of his predecessors, Krauze paid attention mainly to cases of appearance of pigmental birth marks, where their forms and situation compel to suggest the influence of pregnant women. He began with these sharp effects, as terror, fear, anger and so on, showing strong physiological influence expressed first of all in the reaction from the side of vascular and nervous systems. From this Krauze made the conclusion, that "if the brain was strongly alarmed, then small changes took place in the body". Referring to known cases of adult people suddently growing grey under the effect of deep feelings, Krauze suggested that, there were more bases to expect changes of skin color of the fetus under the effect of maternal imagination. If terror or fear can cause small ulcers on lips or erysipelas, then why does it seem incredible to you, that the same phenomena can take place in the body of the fetus, whose structure is so weak and delicate, and its vessels are so numerous and full of juices?"
finished Kiev Ecclesiastical Academy, studied medicine in
Strasburg, where he defended a doctor dissertation about
"human liver" (1755). At Petersburg admiralty hospital, he
read obstetrics and wrote a lot of works and manual books.  
The second part of the family (Ambodic) was written by himself
in connection with accord of patronymic and family (127) .


616


In order to imagine the possibility of this phenomenon, which is shown by the influence of changes in the organism of the mother on the fetus, it is necessary to prove their close relation. On his side Krauze confirmed, "fetus with the uterus represents a single continuous whole". According to this opinion, this is especially related to the nervous system, so that "stimulation of the nerves of the uterus may and must be passed to all the nervous system of the fetus. Distributing this confirmation on psychics, as well as suggesting, that the fetus was capable of psycological manifestations, Krauze deduced the following conclusion: "In the brain of the fetus exists the same condition, which exists in the brain of the mother". All these views are summarized in the concluding paragraph of the dissertation". What was stated so far may be added to all these examples, which instruct, that the fetus body is changed from the mother, if her soul is strongly shocked. Ideas, originating in the mother's brain are "united with fetus brain"; they are stimulated in it more quickly and energetically, than in the soul of the mother herself, as the pulse of arteries and effect of nervous and generally all responses in babies are quicker than in adults. Under the influence of these ideas, the fetus brain gives effect on its body and namely in this way, which corresponds to perceived ideas. Therefore, the fetus's brain produces in the corresponding parts of his body the same things which he himself undergoes".


Directly after the considered work of Krauze, it is necessary to examine the work of Rederer, which is simply entitled "Dissertatis" and also begins with the formulation of the question raised by the Academy: "What is the direct cause for changing the fetus body?..."


(36) "Dictionary" of Maksimovich Ambodic is composed
In order to possibly answer this stated question, Rederer considered that it is necessary to study thoroughly, how the mother's body unites with the embryo, taking into consideration, that the only connection between them is carried out through the placenta. The investigation of this connection led him to the following conclusion. The lumens of the blood vessels of the placenta, directed to the uterus, up till this stage, are so narrow. They do not admit turpentine oil or any other liquid to pass. True anastomoses, connecting the vessels of the uterus and vessels of placenta, do not exist; Vasa hypogastrica which is full, by all means of a waxen mass or any other fluid, does not pass its contents to the umbilical vessels. This was established either in human carcass, or in living cows, ewes, bitches and other animals. If the pregnant animal is fed on roots of Rabiae tinctorum, then the fetus bones will not be stained with the red color. The blood of the fetus differs from the mother's blood in that it is more liquid. Finally, the fetus pulse also differs from the mother's rate.
of two parts: Russian-Latin-French and Latin- Russian, which  
was prefaced with 65 pages of the text, explaining the  
significance of this first Russian terminological dictionary,  
contents of anatomy and physiology and even some information
about these sciences.  


In "the foreward to the dictionary in general",
Subsequently, the fetus lives its independent life, moving its own blood by its forces, beating of its heart, by blood circulation and through placenta, without the help of the maternal blood. Therefore, rest, movement, sleeping, awakening and even life and death of the mother and fetus are not obligatory present in harmony. Later on, Rederer categorically denied the existence of a nervous connection between the placenta and the mother. The comparison of the positions of the birth marks as well as their presence or absence in the mother and baby does not give, according to his opinion, basis to suggest the influence through blood or nervous system. Rederer discussed in detail the question about "mind" of the fetus and passed to the conclusion, that this mind cannot adequately respond to the feeling of the mother. This is stated by the examples of strong shocks in the mother, not accompanied by the appearance of birth marks in the baby, and examples of appearance of birth marks and warts without any connection with the mother's feeling. In the following pages the various deformities of human beings are mentioned, and Rederer reached the conclusion that the birth marks differ from true deformities only quantitatively. They are also disturbances of the normal development. As the deformities frequently affect the internal organs, then in relation to them, the usual supposition about maternal influence loses its significance. The general conclusion of Rederer comes to that the confirmations about the influence of the mother's feeling on the baby are not supported by verified facts, and are but the product of fantasy (105) .
Maksimovich-Ambodic writes: "During collection of the  
words, related to my subject of practice, I have been
collecting bees for more than 10 years. The major reason
was that the Russian words had been collected from various
ancient and recent manuscripts belonging to church and  
civics."


It is difficult to overestimate the significance of this
(27) The effect of vitalistic views of Blumenbackh on some Russian physicians-biologists can be traced up till the 20 's of the 19th century. Thus, in 1825 at Moscow University, the surgeon Nikifor Dmitrievich Lebedev discussed a thesis for the degree of Doctor of Medicine. He later on read at the university "history and literature of medicine". Lebedev dissertation was entitled "About the nature of weightless substances in general and vital powers in particular" (Dissertatio inauguralis physislogica de natura imponderab ilium in genere et de viribus vitalibus in specie, quam...in Universitate caesarea Mosquensi, pro gradu doctoris medicinae. . .elaboravit publiceque defendet chirurgus I-mae classis Nicephorus Lebedev, Mosquae, 1825. 28 p.). Lebedev, referring to Blumenbakh, dwelt on the idea that all vital processes — organic formation and growth as well as movement of already formed parts-possess their own source of a special vital power (55) . The vital power, according to his opinion, is an internal, inherent character in the organic body, which is the cause of life and at the same time seems to be its product (Vis vitalis est interna et proporia organici corpori qualitis, quae vitae causam constituit et simul ejusdem est quasi productum, thesis 9) . Works of similar kind are not characteristic of the common materialistic trend of Russian biological and medical sciences. Therefore, the dissertation of Lebedev served as a reference and mainly as an illustration of this negative influence, which was shown by the idealistic German philosophy and some representatives of Russian sciences especially the naturalists who worked under its direct influence (106) .
work as a result of the great quantity of terms created, which
were absent in the Russian language. This significance is  
not diminished by the bulkness of some terms which are not  
contained in scientific language together with others which
are archaic and have disappeared from the language in the  
process of its evolution.  


The historian of national medicine Ya. A. Chistovich
(28) Izef Gotlib Kelreiter (1733-1806), botanist and zoologist, worked from 1756 to 1761 as a junior scientific assistant at Petersburg Academy of Sciences, with which he kept in close contact after returning back to Germany and till the end of his life. In the period, from 1758 to 1811, 15 botanical and more than 20 zoological works of Kelreiter were published in Russian scientific and scientific popular editions (Novi Commentarii Acad. Scient. petropolitanae, Acta Acad. Scient. petropol. Nova Acta Ac. Sc. Petropl., Trudy Volnogo Ekonomicheskogo Obshchestva and in the journal "Sochineniya,K poize i uveceleniyu sluzhashchie") . His scientific fame is connected mainly with the study of reproduction and hybridization in plants (see Ioz. Kelreiter" Study about sex and hybridization in the plant", the editor with a biological essay was prof. E. V. Wolff, 1840).
drew a special attention to the significance of "the dictionary" of Maksimovich-Ambodic. He noted the unfair relation
to Maksimovich on the part of V. M. Richter, who "did not
offer him a single line in his biographical History of
Medicine in Russia". Fortunately, the voice of this light
word was fair and, in defiance of the partial historian,  
preserved the name of Maksimovich from undeserved oblivion"
(Ya. Chistovich. First obstetrician schools in Russia
(1754-1785. Essays from history of Russian medical
institutions of the 18th century. SPb., 1870, p. 199) (131).  


(37) Khristian Elias Genrikh Knakshtedt was born in
Braunshveig in 1749, studied surgery in Bryunn and in 1786
travelled to Petersburg, where he was professor of anatomy
and surgery at Kalinkinsky hospital. In 1790, due to the
work "Beschreibung der trockenen Knochen des menschlichen
Korpers" (SPb., 1791) the medical college awarded him the
degree of Doctor of Medicine. Knakshtedt died in 1799. In
addition to the mentioned works, he also published "Descriptio
praeparatorum maximam partem osteologicorum rarissimorum"


617
The work of Kelreiter about the irritability in the plant, to which Wolff referred, was called: "Nouvelles observations et experiences sur l'irritabilite des etamines de l'epine vinette (Berberis vulgaris)", Traduit de l'allemand par M. I 1 adjoint Sewergin, Nova Acta Acad. Scient. Petropol., 4, 1790 (German original was received in Petersburg in 1788) (112).


(29) The point of view of A. E. Gaisinovich is similar to the presentation mentioned here about the outlook of Wolff (see Wolff "Theory of conception") . It is possible to combine with it the confirmation, that the studies on preformation and epigenesis do not always correspond to the demarcation between idealism and materialism in biology. The following serves as an evidence, that preformists were also idealists (Leibnitz, Haller, Bonnet) and mechanical materialists (for example, Lamettri) ; equally as epigenetics, were either idealists (Aristotle Harvei) ,


or mechanical materialists (Dekart, Maupertuis, Byuffon and Didar3t).A. E. Gaisinovich included Wolff as well in the latter category. His "absolute epigenesis" was grouped together with Gaisinovich in the mechanical materialism. For its complete verification, this statement could be desired as only a less categorical expression. In the evaluation of Wolff's outlook, it is impossible, apparently, to deny absolutely his fluctuations between materialism and idealism; these fluctuations were an unavoidable originality even by the most prominent thinkers of all historical epochs, preceding the formation of successive system, i.e. dialectic system and materialism (119) .


(Braunschweig, 1785) and Latin-German terminological medical  
(30) Semen Gerasimovich Zybelin (1735-1802), after finishing the study in Moscow Ecclesiastical Academy, joined Moscow University till the opening there of a medical faculty. After he finished the course, he was sent to study medicine abroad. At Leiden, Zybelin defended a doctor's dissertation in 1764. After returning back to Moscow he read courses of anatomy, physiology, chemistry, pharmacology and therapy as a professor in the mediqal faculty. In 1784, he was elected member of the Academy of Science (121) .
dictionary "Erklarung lateinischen Worter, welche zur
Geliederungslehre Physiologie; Wundarzneywissenschaft und...
Ordnung" (Braunschw., 1784, 2nd edition, SPb., 1788). In
the title page of the latter he called himself Russian surgeon
and ordinary teacher of studies about bones and all their
diseases at the imperial surgical school in Petersburg.  


To the work of Knakshtedt mentioned in the text
(31) Prince Dmitrii Alekseevich Golitsyn (1731-1803), a prominent Russian diplomat, was a former ambassador in Holland and France, friend of Diderot and Galvetsy, known as author of many physical works (122) .
"Anatomical description of the monster" an invitation to
Medico-surgical school is addressed to "all famous persons
and members of medico- surgical sciences" to attend the meeting
"near Kalinkinsky bridge at Ekateringofskaya and listen to
"some works and speeches by some teachers and students.
The meeting was proposed to take place on January 7 at
10 a.m. (131) .  


(38) Petr Andr^evich Zagorskii (1761-1845) is the one
(32) Dzh. Needham in his "History of Embryology" prefaced the list of the literature sources used by him with a list of works, which he could not obtain. As "less important works on the history of embryology" Needham also mentioned Bezeke's book. It is difficult to decide, on which basis the English historian of embryology considered Bezeke's book less important, if we take into consideration that according to his confession, he never saw it (122) .
who founded the first Russian anatomical school . After
teaching at the Cheringovsky college he went to the hospital
school in Petersburg. After he finished there, he worked for
three years as prosecutor Petersburg medico-surgical school
with professor N. P. Karpinsky. From 1799 to 1833, he was
junior assistant, and then professor in medico-surgical
school, and from 1805 up to the end of his life he was a
member of the Academy of Sciences. Zagorskii organized
excellent anatomical museums at the medico-surgical school
and Academy of Sciences, published the first Russian manual
book on anatomy and a great number of works on anatomy,  
teratology and different medical topics.  


(38a) The same opinion on the origin of monsters was
(33) Two extracts compose the contents of the already mentioned book of 1797 (in it, as stated, were present an essay on the history of a hypothesis about the conception and development of animals and in addition, "History of the origin of division of the natural bodies into three kingdoms") The third extract appeared in the form of a separate small volume in the year of the author's death (J. M. G. Bezeke. Allgemeine Geschichte der Naturgeschichte in dem Zeitraume von Erschaffung der Welt bis auf das Jahr N. C. G. 1791. Mitau, 1802, XXXII±154S) . For more details about the works of Bezeke see the article by the author of the present book in "Trudy instituta istoriii estestvoznaniya i tekhniki, V.IV, 1955" (Works of the Institute of History of Natural Sciences and Technique) (123).
also supported by the academician N. Ya. Ozeretskovskii, who
informed the Academy on April 25, 17991 about two cases of  
double monsters in the preparations of the academic museum of  
natural history. One of these cases was united twins (union
in the region of the upper part of the chest, both partners


(34) Matvei Khristianovich Peken was born in Petersburg, he studied medicine in Ien, where he obtained the degree of Doctor of Medicine (according to other data-- in Gettingen) . When he returned back to Russia he worked as the admiralty doctor, read a course of obstetrics in Petersburg hospitals. From 1793, he travelled to Moscow, where he read pathology and organized the first therapeutic clinic with ten beds (124)


(35) Nestor Maksimovich Maksimovich-Ambodic (1744-1812) finished Kiev Ecclesiastical Academy, studied medicine in Strasburg, where he defended a doctor dissertation about "human liver" (1755). At Petersburg admiralty hospital, he read obstetrics and wrote a lot of works and manual books. The second part of the family (Ambodic) was written by himself in connection with accord of patronymic and family (127) .


1. N. Ozeretskovskii. De doubus foetibus humanis,
(36) "Dictionary" of Maksimovich Ambodic is composed of two parts: Russian-Latin-French and Latin- Russian, which was prefaced with 65 pages of the text, explaining the significance of this first Russian terminological dictionary, contents of anatomy and physiology and even some information about these sciences.


monstrosis. Nova Acta Acad. Sc. imp. Petropol., 14.
In "the foreward to the dictionary in general", Maksimovich-Ambodic writes: "During collection of the words, related to my subject of practice, I have been collecting bees for more than 10 years. The major reason was that the Russian words had been collected from various ancient and recent manuscripts belonging to church and civics."
1805, p. 367-372. The article of Ozeretskovskii was
published in the same volume of Nova Acta Academiae
Petropolitanae , where the above mentioned report of
Zagorskii was also published.  


618
It is difficult to overestimate the significance of this work as a result of the great quantity of terms created, which were absent in the Russian language. This significance is not diminished by the bulkness of some terms which are not contained in scientific language together with others which are archaic and have disappeared from the language in the process of its evolution.


The historian of national medicine Ya. A. Chistovich drew a special attention to the significance of "the dictionary" of Maksimovich-Ambodic. He noted the unfair relation to Maksimovich on the part of V. M. Richter, who "did not offer him a single line in his biographical History of Medicine in Russia". Fortunately, the voice of this light word was fair and, in defiance of the partial historian, preserved the name of Maksimovich from undeserved oblivion" (Ya. Chistovich. First obstetrician schools in Russia (1754-1785. Essays from history of Russian medical institutions of the 18th century. SPb., 1870, p. 199) (131).


(37) Khristian Elias Genrikh Knakshtedt was born in Braunshveig in 1749, studied surgery in Bryunn and in 1786 travelled to Petersburg, where he was professor of anatomy and surgery at Kalinkinsky hospital. In 1790, due to the work "Beschreibung der trockenen Knochen des menschlichen Korpers" (SPb., 1791) the medical college awarded him the degree of Doctor of Medicine. Knakshtedt died in 1799. In addition to the mentioned works, he also published "Descriptio praeparatorum maximam partem osteologicorum rarissimorum"


were completely formulated) . The second case was the
doubling of the head end, beginning from the girdle region;
the monster had three correctly- formulated hands, two normal
and one underdeveloped legs. The description of the monsters
(the first case was illustrated by two excellently-engraved
figures, representing the twins from the front and from
behind) was concluded with a brief account about the reason
for the monsters appearance. By comparison of the described
cases, Ozeretskovskii made the conclusion that monsters
can be very variable and that each monster, must possess
its natural cause. "The physiologists — Ozeretskovskii
wrote — must explain these causes and find the specific
original source of these monsters — whether their origin is
due to the union of two embryos, or from strengthened,
weakened or incomplete development of parts of the body"
(p. 371).


In another article, whose contents were received by the
(Braunschweig, 1785) and Latin-German terminological medical dictionary "Erklarung lateinischen Worter, welche zur Geliederungslehre Physiologie; Wundarzneywissenschaft und... Ordnung" (Braunschw., 1784, 2nd edition, SPb., 1788). In the title page of the latter he called himself Russian surgeon and ordinary teacher of studies about bones and all their diseases at the imperial surgical school in Petersburg.
Academy one year before (on October 25, 1798)1, a description
was given for non-hatched hen's egg with an opening in the
shell; through this opening the end of a blood vessel passed.  
When the egg was opened, in addition to yolk and egg white,  
a pear-shaped body was found in it, which was full of clotted
blood. Ozeretskovskii considered the described content in
the egg as a polyp of the oviduct, torn at the time of yolk
passing, falling in the egg together with the white. In  
connection with this, he assumed, that the presence of similar
kinds of strange bodies in the eggs of birds may be the cause
of appearance of monsters, as the mechanical pressure on the
delicate parts of the developed embryo inevitably leads to
their deformation. During this, Ozeretskovskii referred to
cases of development in double-yolk eggs of doubled embryos
which — as a rule — are monsters. During more or less normal
development of these twins, they can unite with each other.
"Some years before" Ozeretskovskii wrote "we saw here, in
Petersburg, doubled chickens, hatched from one egg, provided
with all organs and united at the backs; when one of them
stood on the legs, the other lied on it on the back with legs
upwards in the most unnatural position. It is clear that,


To the work of Knakshtedt mentioned in the text "Anatomical description of the monster" an invitation to Medico-surgical school is addressed to "all famous persons and members of medico- surgical sciences" to attend the meeting "near Kalinkinsky bridge at Ekateringofskaya and listen to "some works and speeches by some teachers and students. The meeting was proposed to take place on January 7 at 10 a.m. (131) .


(38) Petr Andr^evich Zagorskii (1761-1845) is the one who founded the first Russian anatomical school . After teaching at the Cheringovsky college he went to the hospital school in Petersburg. After he finished there, he worked for three years as prosecutor Petersburg medico-surgical school with professor N. P. Karpinsky. From 1799 to 1833, he was junior assistant, and then professor in medico-surgical school, and from 1805 up to the end of his life he was a member of the Academy of Sciences. Zagorskii organized excellent anatomical museums at the medico-surgical school and Academy of Sciences, published the first Russian manual book on anatomy and a great number of works on anatomy, teratology and different medical topics.


1. N. Ozeretskovskii. De ovo perforata. There also,  
(38a) The same opinion on the origin of monsters was also supported by the academician N. Ya. Ozeretskovskii, who informed the Academy on April 25, 17991 about two cases of double monsters in the preparations of the academic museum of natural history. One of these cases was united twins (union in the region of the upper part of the chest, both partners were completely formulated) . The second case was the doubling of the head end, beginning from the girdle region; the monster had three correctly- formulated hands, two normal and one underdeveloped legs. The description of the monsters (the first case was illustrated by two excellently-engraved figures, representing the twins from the front and from behind) was concluded with a brief account about the reason for the monsters appearance. By comparison of the described cases, Ozeretskovskii made the conclusion that monsters can be very variable and that each monster, must possess its natural cause. "The physiologists — Ozeretskovskii wrote — must explain these causes and find the specific original source of these monsters — whether their origin is due to the union of two embryos, or from strengthened, weakened or incomplete development of parts of the body" (p. 371).
12.1801, pp. 364-368.  






619
1. N. Ozeretskovskii. De doubus foetibus humanis, monstrosis. Nova Acta Acad. Sc. imp. Petropol., 14. 1805, p. 367-372. The article of Ozeretskovskii was published in the same volume of Nova Acta Academiae Petropolitanae , where the above mentioned report of Zagorskii was also published.






only the narrowness of the shell was the cause that the twin  
In another article, whose contents were received by the Academy one year before (on October 25, 1798)1, a description was given for non-hatched hen's egg with an opening in the shell; through this opening the end of a blood vessel passed. When the egg was opened, in addition to yolk and egg white, a pear-shaped body was found in it, which was full of clotted blood. Ozeretskovskii considered the described content in the egg as a polyp of the oviduct, torn at the time of yolk passing, falling in the egg together with the white. In connection with this, he assumed, that the presence of similar kinds of strange bodies in the eggs of birds may be the cause of appearance of monsters, as the mechanical pressure on the delicate parts of the developed embryo inevitably leads to their deformation. During this, Ozeretskovskii referred to cases of development in double-yolk eggs of doubled embryos which — as a rule — are monsters. During more or less normal development of these twins, they can unite with each other. "Some years before" Ozeretskovskii wrote "we saw here, in Petersburg, doubled chickens, hatched from one egg, provided with all organs and united at the backs; when one of them stood on the legs, the other lied on it on the back with legs upwards in the most unnatural position. It is clear that, only the narrowness of the shell was the cause that the twin chickens, which were pressed to each other, united by the backs, similar to the united apples, which, beginning from the moment of flowering, closely adjoined each other".
chickens, which were pressed to each other, united by the  
backs, similar to the united apples, which, beginning from  
the moment of flowering, closely adjoined each other".  


Both mentioned articles of Ozeretskovskii witness that
the Russian academician considered the deformity a result of
the caused changes of normal development by external influences, i.e. explained their appearance epigenetically (134).


(39) K. Fr. Kielmeyer (1765-1844), is a famous German
naturalist, held in great respect by the contemporaries
although he hardly left published works. The most famous
work of Kielmeyer was his speech "About the relation of
organic powers between each other" (1793) . Kielmeyer
particularly suggested the idea of powers, inherent within
the living beings — irritability, sensibility and reproduction. Their combination in the form of ascending and
descending rows corresponds to the stage of the individual
development as well as the stage of development of all the
organic world, in which, according to Kielmeyer, "the plan
of nature" is found. By comparison of the stages of development with these rows, in which adult forms may exist,
Kielmeyer made a conclusion, stated by him in a conversation
with Gete (1797), that the higher organisms pass many stages
in the process of embryonic development, in which they become
lower. Shelling attached a very great importance to the
ideas of Kielmeyer considering, that they usher in a new
epoch in science. In fact, he found later on that before
Kielmeyer a similar idea was stated by I. G. Gerder (17441863), who was a publicist, poet and philosopher. The works
of Gerder, in particular, had a relation to the question
discussed — "The ideas of the philosophy of the humanity
history", were highly evaluated by Gete. They were popular
in Russia as well and had a known effect on Karamzin,
Shevryev and Maksimovich (143) .


(40) Prince A. P. Baryatinskii a personality of the
1. N. Ozeretskovskii. De ovo perforata. There also, 12.1801, pp. 364-368.
Southern Secret Society and active propagandist became,  
towards the end of 1825, the chief of Kilchinskaya board


(in Tul'chin the staff of the 2nd army was present, in which
Pestel and some other members of Southern Secret Society
served) (148) .


Both mentioned articles of Ozeretskovskii witness that the Russian academician considered the deformity a result of the caused changes of normal development by external influences, i.e. explained their appearance epigenetically (134).


(39) K. Fr. Kielmeyer (1765-1844), is a famous German naturalist, held in great respect by the contemporaries although he hardly left published works. The most famous work of Kielmeyer was his speech "About the relation of organic powers between each other" (1793) . Kielmeyer particularly suggested the idea of powers, inherent within the living beings — irritability, sensibility and reproduction. Their combination in the form of ascending and descending rows corresponds to the stage of the individual development as well as the stage of development of all the organic world, in which, according to Kielmeyer, "the plan of nature" is found. By comparison of the stages of development with these rows, in which adult forms may exist, Kielmeyer made a conclusion, stated by him in a conversation with Gete (1797), that the higher organisms pass many stages in the process of embryonic development, in which they become lower. Shelling attached a very great importance to the ideas of Kielmeyer considering, that they usher in a new epoch in science. In fact, he found later on that before Kielmeyer a similar idea was stated by I. G. Gerder (17441863), who was a publicist, poet and philosopher. The works of Gerder, in particular, had a relation to the question discussed — "The ideas of the philosophy of the humanity history", were highly evaluated by Gete. They were popular in Russia as well and had a known effect on Karamzin, Shevryev and Maksimovich (143) .


620
(40) Prince A. P. Baryatinskii a personality of the Southern Secret Society and active propagandist became, towards the end of 1825, the chief of Kilchinskaya board (in Tul'chin the staff of the 2nd army was present, in which Pestel and some other members of Southern Secret Society served) (148) .




(41) Efrem Osipovich Mukhin (1766-1850) studied at the Kharkovsky College and Elisavetgradskaya Hospital School, and then at Moscow University. He was a junior assistant at the department of pathology and therapy at Moscow Medical School, and later on a professor at the university. He wrote a great number of manual books and special works (162) .


(41) Efrem Osipovich Mukhin (1766-1850) studied at  
(42) Vilglin Michailovich Rikhter (1767-1822) was born in Moscow, finished Moscow University and published a doctor's dissertation in Erlangen. From 1790 he was a professor of obstetrics at Moscow University. Rikhter is the author of project "Practical obstetrical institute" and many manual books (162) .


the Kharkovsky College and Elisavetgradskaya Hospital School,  
(43) Ivan Fedorovich Wenssowitsch (1769-1811) studied at the Kharkovsky college and Moscow University secondary school, from where he joined the university, where he successively joined the faculties of philosophy, law and medicine. He finished the university and became a candidate of medicine. In 1803, he defended a doctor's dissertation in Moscow. From 1805 he was a professor of anatomy, physiology and forensic medicine. He published "journal medico-physical He died from tuberculosis at the age of 42 (163) .
and then at Moscow University. He was a junior assistant at
the department of pathology and therapy at Moscow Medical
School, and later on a professor at the university. He
wrote a great number of manual books and special works (162) .  


(42) Vilglin Michailovich Rikhter (1767-1822) was
(44) Yakov Kuzmich Kaidanov (1779-1885), from Kiev Ecclesiastical Academy, studied in the Petersburg medicosurgical school. After finishing there, he was sent to Vienna to study veterinary medicine. After his return back, he became a junior assistant at the Medico-surgical Academy. From 1809, he became a
born in Moscow, finished Moscow University and published a
doctor's dissertation in Erlangen. From 1790 he was a  
professor of obstetrics at Moscow University. Rikhter is
the author of project "Practical obstetrical institute"
and many manual books (162) .  


(43) Ivan Fedorovich Wenssowitsch (1769-1811) studied
at the Kharkovsky college and Moscow University secondary
school, from where he joined the university, where he
successively joined the faculties of philosophy, law and
medicine. He finished the university and became a candidate
of medicine. In 1803, he defended a doctor's dissertation in
Moscow. From 1805 he was a professor of anatomy, physiology
and forensic medicine. He published "journal medico-physical
He died from tuberculosis at the age of 42 (163) .


(44) Yakov Kuzmich Kaidanov (1779-1885), from Kiev
{{Blyakher1955 footer}}
Ecclesiastical Academy, studied in the Petersburg medicosurgical school. After finishing there, he was sent to
Vienna to study veterinary medicine. After his return back,
he became a junior assistant at the Medico-surgical Academy.
From 1809, he became a

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Blyakher L. History of embryology in Russia from the middle of the eighteenth to the middle of the nineteenth century (istoryia embriologii v Rossii s serediny XVIII do serediny XIX veka) (1955) Academy of Sciences USSR. Institute of the History of Science and Technology. Translation Smithsonian Institution (1982).

   Historic Russian Embryology 1955: 1. Beginning of Embryological Investigations Lomonosov's Epoch | 2. Preformation or New Formation? | 3. Kaspar Friedrich Wolff - Theory of Epigenesis | 4. Wolff: "Theory Of Generation" | 5. Wolff: "Formation of the Intestine" | 6. Wolff's Teratological Works | 7. Wolff: "On the Special Essential Tower" | 8. Ideology of Wolff | Chapter 9. Theory of Epigenesis End of 18th Century | 10. Embryology in the Struggle of Russian Empirical Science Against Naturphilosophie | 11. Louis Tredern - Forgotten Embryologist Beginning of 19th Century | 12. Embryonic Membranes of Mammals - Ludwig Heinrich Bojanus | 13. Embryonic Layers - Kh. I. Pander | 14. Karl Maksimovich Baer | 15. Baer's - De Ovi Mammalium Et Hominis Genesi | 16. Baer's Ober Entw I Cklungsgesch I Chte Der Thiere | 17. Baer Part 1 - Chicken Development | 18. Baer Part 2 - History of Chicken Development | 19. Baer Vol 2 | 20. Third Part of the Bird Egg and Embryo Development | 21. Third Part - Development of Reptiles, Mammals, and Animals Deprived of Amnion and Yolk Sac | 22. Fourth Part - Development of Man | 23. Baer's Teratological Works and Embryological Reports in Petersburg | Chapter 24. Baer's Theoretical Views | 25. Invertebrate Embryology - A. Grube, A. D. Nordmann, N. A. Warnek, and A. Krohn
Online Editor 
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This historic textbook by Bliakher translated from Russian, describes historic embryology in Russia between 1750 - 1850.



Publishing House of the Academy of Science USSR

Moscow 1955

Translated from Russian

Translated and Edited by:

Dr. Hosni Ibrahim Youssef # Faculty of Veterinary Medicine Cairo University

Dr. Boulos Abdel Malek

Head of Veterinary Research Division

NAMRU-3, Cairo

Arab Republic of Egypt

Published for

The Smithsonian Institution and the National Science Foundation, Washington, D.C, by The Al Ahram Center for Scientific Translations 1982


Published for

The Smithsonian Institution and the National Science Foundation, Washington, D.C by The Al Ahram Center for Scientific Translations (1982)


Also available online Internet Archive


Historic Embryology Textbooks

Historic Disclaimer - information about historic embryology pages 
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Pages where the terms "Historic" (textbooks, papers, people, recommendations) appear on this site, and sections within pages where this disclaimer appears, indicate that the content and scientific understanding are specific to the time of publication. This means that while some scientific descriptions are still accurate, the terminology and interpretation of the developmental mechanisms reflect the understanding at the time of original publication and those of the preceding periods, these terms, interpretations and recommendations may not reflect our current scientific understanding.     (More? Embryology History | Historic Embryology Papers)

Academy of Sciences USSR

Institute of the History of Science and Technology

A. N. Severtsov Institute of Animal Morphology


L. Ya. Blyakher

History of Embryology in Russia from the Middle of the Eighteenth to the Middle of the Nineteenth Century

(Istoryia Embriologii v Rossii s serediny XVIII do serediny XIX veka)

Institute of the History of Science and Technology

A. N. Severtsov Institute of Animal Morphology

L. Ya Blyakher

Editor: G. A. Shmidt

With an Introduction by Jane Maienschein

Publishing House of the Academy of Science USSR

Moscow 1955

Contents

Introduction by Jane Maienschein

Preface

  1. The Beginning of Embryological Investigations in Russia in Lomonosov's Epoch
  2. Preformation or New Formation?
  3. Kaspar Friedrich Wolff and Substantiation of the Theory of Epigenesis
  4. The "Theory Of Generation" of K. F. Wolff
  5. Wolff's Treatise "On the Formation of the Intestine"
  6. Wolff's Teratological Works
  7. The Work of Wolff: "On the Special Essential Tower"
  8. The Ideology of Wolff
  9. The Theory of Epigenesis is in Russia at the End of the 18th Century
  10. The Development of Embryology in the Epoch of the Struggle of Russian Empirical Science Against Naturphilosophie
  11. Louis Tredern - The Forgotten Embryologist of the Beginning of the Nineteenth Century
  12. The Study of the Embryonic Membranes of Mammals - The Investigations of Ludwig Heinrich Bojanus
  13. The Discovery of Embryonic Layers - The Dissertation of Kh. I. Pander
  14. Outline of the Life and Scientific Activities of Karl Maksimovich Baer
  15. Baer's Treatise De Ovi Mammalium Et Hominis Genesi
  16. Appearance and Significance of Baer's Main Work Ober Entw I Cklungsgesch I Chte Der Thiere
  17. First Part of Uber Entw I Cklungsgesch I Chte - Development Of The Chicken in the Egg
    1. First Period of Chick Development
    2. The Second Period of Chick Development
    3. The Third Period of Chick Development
  18. Second Part Of Uber Entwicklungsgeschichte - Scholia and Corollaries to the History of Chicken Development in the Egg
  19. Theoretical Introduction to the Second Volume of Uber Entwicklungsgeschichte
  20. Third Part of Uber Entw I C Klungsgesch I Chte - Development of the Bird Egg and Embryo
  21. Third Part of Flber Entwi Cklungsgesch I Chte Continued - Development of Reptiles, Mammals, and Animals Deprived of Amnion and Yolk Sac
  22. Fourth Part of Uber Entwicklungsgeschichte - Studies on the Development of Man
  23. Baer's Teratological Works and his Embryological Reports, Related to the Period of his Work in Petersburg
  24. On the Question of Baer's Theoretical Views
  25. Investigations on Invertebrate Embryology - Work of A. Grube, A. D. Nordmann, N. A. Warnek, and A. Krohn


Comments

Illustrations To Blyakher, History Of Embryology In Russia

Introduction by Jane Maienschein

General

When Dr. Robert Multhauf asked me if I would consider editing this translation of Blyakher's volume, he warned that this was part of what seemed to him a most unusual scholarly project. Thanks to a somewhat mysterious and complicated government exchange program, the Smithsonian Institution and the National Science Foundation had been charged with overseeing the translation into English of several foreign language texts in the history of science. Upon the recommendation of experts, the volumes chosen included two by L. Blyakher, a Russian biologist. In particular, these Russian volumes, including THE HISTORY OF THE INHERITANCE OF ACQUIRED CHARACTERISTICS, edited by Frederick Churchill, and this one, were thought to present a valuable exposure to a Russian point of view in the history of science and to detail important episodes of Russian scientific history. Therefore, the translation began.

Following someone's recommendation, this particular volume went to Egypt to be translated by an anonymous translator. I admire the translator's patience in working through the detailed embryological descriptions. Unfortunately, however, the style of the English translation was infelicitous at best, and the translator evidently had trouble with proper names, German references, and embryological terms. My task, then, became to turn the prose into an acceptable style, to correct the names and terms to conform with standard English usage, and to check the references.

The fact that I do not read Russian, except for wordby-word translation with a dictionary, could have posed a fatal problem. But fortunately, the Dickinson College Library and work study office generously donated the services of work study student Lauri Wiener, who reads Russian and possesses the requisite active curiosity and healthy sense of humor. Together, Lauri and I checked the questionable phrases as well as a random sample of other passages to determine the accuracy of the translation. Except for some of the discussion


of German philosophy and a few embiyological descriptions, the translation appeared to us to be accurate. The fact that Blyakher's style is straightforward and essentially descriptive undoubtedly helped, since the translator could thus provide a rather literal translation without losing the content or warping the style significantly. The fact that many of the embryological terms had simply been transliterated from Latin or German into Russian and then back in accordance with standard international scientific terminology to make this translation, meant that the terms usually remained recognizable. Thus, although I make no pretense of certifying the precision of every detail of this translation, to my knowledge it is reasonably accurate at all points and represents Blyakher's content and style fairly closely. I very much appreciate Lauri Wiener's help in verifying and improving the translation.

Identifying some of the intended proper names and dates required a bit of detective work. Double transliteration or translation into Russian and then into English created much more trouble with some of the names than did translation of embryological terms. Names such as Isidore Geoffroy Saint-Hilaire or Cuvier produced mysterious and occasionally hilarious results, as the former became Izedor Zhefwar Tzent IJer, and the latter Kyuve. Joseph Needham became Nidzhem, Xeeuwenhoek became Lev'nhk, and so on. As might be expected, the more obscure names created the greatest difficulties, but with the help of the extraordinarily helpful and competent Dickinson College Interlibrary Loan staff, I managed to track down all but a couple of minor Russian figures to check spellings and dates. In questionable cases, I have used spellings from the Library of Congress National Union Catalog. And some names are left in Russian style, such as Karl Maksimovitch Baer (alias Karl Ernst von Baer, of course) to reflect the importance of Blyakher's claim that these men are essentially Russians. In this case, Karl Maksimovitch Baer is closer to the man's given name when he was born in Estonia.

References to published and unpublished materials provided even more trouble in some cases, though here, too, I was able to check and correct all except a few Russian references. A project of this type and magnitude naturally encourages some errors to creep in, so I expect that there


may be some imperfections in citations. Nonetheless, with the exception of some of the Russian articles, I have been able to verify dates, page numbers, and other significant reference data. Readers with access to a superior library should be able to locate most of the material Blyakher cites, though some of the unpublished Russian materials may well prove inaccessible as they did to me.

My other task in editing this volume lay in making the descriptive chapters on von Baer's UBER ENTWICKELUNGSGESCHICHTE useful, since Blyakher 's page number references to the Russian translation of von Baer's work would obviously not be particularly relevant for most readers of this English translation. Therefore, for the passages Blyakher quotes or cites, I identified page numbers of the German original edition, which has been reprinted recently. And where necessary I checked, corrected, and added the section references to UBER ENTWICKELUNGSGESCHICHTE. The references are thus: (volume, section, German page number) or (volume, section, Russian page number (German page number)). In addition, I corrected translation of the quotations where necessary to accord with more common English versions .

For other references to or quotations from German or English works, I made necessary corrections and substituted standard English versions where available since some passages had been distorted in the double translation.

At this point, the first round of editing was complete. Here Rosemary Regan enters the story. Ms. Regan, a marvelously competent and intelligent assistant to Dr. Multhauf, helped with typing some of the longer and messier chapters; she corrected errors in the entire draft, and she used her knowledge of editing and the history of science to polish details of style and terminology. I thank Rosemary for her considerable help and both Rosemary and Robert Multhauf for their continued encouragement and good humor.

With these acknowledgements and with the above caveats, I feel assured that this descriptive volume should be accurate and usable. Editing this has proven to be an unusual project, as Dr. Multhauf warned, but I feel, as he has felt, that the translated and edited volume can prove useful, as indicated in more detail below.

Outline of Blyakher's Work

In fact, this book represents only half of Blyakher's HISTORY OF EMBRYOLOGY IN RUSSIA, covering only the mid-eighteenth to the mid-nineteenth century. A brief discussion of the second volume, covering the mid-nineteenth to the mid-twentieth century, appears in a review by Charles Bodemer in ISIS. While that second volume describes material less well-known to western historians of science and while it might therefore seem more valuable, this volume is intriguing in part precisely because it deals with apparently familiar figures and works from a different perspective which is distinctly, though on the whole not zealously, patriotically Russian. Blyakher claims that those recognized great embryologists, Kaspar Friedrich Wolff, Khristian Pander, and Karl Maksimovich Baer - heretofore considered German embryologists - were in fact distinctly Russian, and that their Russian connections define their scientific characters and help explain their successes in important ways. This Russian viewpoint is the first of the book's two major offerings.


The second lies in the description and catalog of essentially inaccessible works and the compiling of several descriptions into a single narrative index of sorts. Blyakher discusses works of major embryologists which can be located only with difficulty. For example, even though they appeared in German (which can be read by more English speakers than Russian can), many of the papers cited appeared in some of the little-circulated publications of the St. Petersburg Academy of Science. Thus, Blyakher provides new descriptions which he combines with discussion of other both major wellknown and relatively rare sources. Since many of these sources are little known, little circulated, inaccessible, or difficult to read because of the archaic scientific detail or descriptive style, Blyakher has performed a valuable service by describing them.


I will outline the chapters briefly as a guide to Blyakher's work, since this is a descriptive study which could use some index, and its indices have not been translated into English for logistical reasons.


In the INTRODUCTION, Blyakher explains that he will give "a detailed account of the history of Russian embryological investigations" to provide "exhaustive evidence (for) the frequently repeated claim that Russia is the fatherland of embryology as a science, that it developed from Russian soil and became one of the most important foundations of the evolutionary and historical view of the organic world." Embryology - meaning Russian embryology, of course - fell into three distinct periods, according to Blyakher: that of establishing epigenesis and making embryology possible as a science (Wolff) , that of discovering the embryonic layers and establishing the prerequisites for comparative embryological development (Pander and Baer) , and that of evolutionary embryological development (Alexander Kovalevsky and I.I. Mechnikov) . The first two periods form the focus of this volume, while the third is subject of his second volume. The following chapters amass "evidence" for his claim for Russian fatherhood primarily by describing the many accomplishments of native (and adopted) Russians and by showing how these actually were in some essential way Russian accomplishments.


CHAPTER 1 considers the early period, beginning with the late seventeenth and the early eighteenth century, the time of Peter I's reorganization in Russia. Peter wanted to encourage native-born and trained scientists, Blyakher tells us, and so the ruler established a significant teratological and embryological collection in the Kunstkamera to support native medical studies. Peter I initiated Russian interest in embryology, according to the author. And Mikhail Lomonosov stimulated serious embryological studies, drawing on those teratological and embryological collections. Lomonosov, like Wolff and other followers, began the modern era of experimentation, materialism, empiricism, and historical explanation, Blyakher asserts, without fully explaining what he means by each of those recurring terms. Despite his infrequent lapses into enthusiastic excesses, Blyakher convincingly establishes that there was early embryological interest in Russia, which has not been widely studied, and that historians of science should therefore explore the subject more seriously.


CHAPTER 2 discusses the preformation and new formation (alias epigenesis) exchanges. This chapter offers few new insights into these debates, but the reader should recall that Blyakher was writing in the early 1950 ! s and that his Russian audience was likely unfamiliar (or only recently familiar) with material which a western audience might find much more familiar. Blyakher' s listing of partisans on either side and his discussion of the issues here and in later chapters are essentially clear and potentially useful even if not profound.


CHAPTER 3 introduces that great adopted Russian, Kaspar Friedrich Wolff, who then provides the subject of Chapters 5-8 as well. Wolff deserves more credit than he has received (by 1955, remember), Blyakher asserts; everyone from Russian historians to German historians to Wolff's contemporaries have reportedly been consistent in their underestimation of Wolff's significance. Here, Blyakher becomes a bit zealous in his efforts to make Russian everyone and everything which seems good or important. He faults the Russian historian and biographer Boris Evgen'evic Raikov, for example, for suggesting that Wolff felt ideologically isolated after his move to Russia in 1767. Not the case, Blyakher insists. ". . .in Germany Wolff was not evaluated as a first class investigator and advanced thinker. This forced him to move to Russia, and therefore Germany does not have the right to claim Wolff's glory."


While Blyakher 's claim is silly as stated and while it might seem exaggerated and annoying to the modern historian of science, it may also reveal valuable insight. It would be well for historians of biology to recall that Wolff was little known in Germany; that he did move from Germany to Russia in 1767, albeit after much of his major embryological work had been completed and published; that his biographers have found the reasons for his move unclear; but that St. Petersburg did offer important collections of embryological and teratological specimens and that Wolff seems to have used them to advantage. Thus, perhaps St. Petersburg did offer an especially congenial environment for an embryologist who was an epigenesist, and perhaps the reasons should be better examined.


CHAPTER 4 provides a useful outline of Wolff's dissertation - both the original Latin of 1759 and the German, more "popular" version of 1764. The Latin criticized earlier epigenetic suggestions and reflected a great deal of respect for Haller, but Blyakher claims that Wolff appealed to Haller only because he sought the latter 's support and that Wolff consistently rejected any tendency toward Haller's preformationist views. In the German, Wolff provided an epigenetic discussion of development and expressed opposition to rigorously mechanical understandings of vital phenomena. Blyakher' s description of Wolff's work is valuable, but the reader should be aware that Blyakher has probably had to strain the data here more than elsewhere to support his thesis about Russian priorities in embryology and his view that Wolff was one of the Russian "good guys" on the progressive path to modern scientific embryology. Again, the reader should recall that this was published in 1955, just shortly after other Russian publications of histories of embryology and translated embryological works.

CHAPTER 5 remains somewhat more descriptive, providing a valuable discussion of Wolff's relatively rarely read ON THE FORMATION OF THE INTESTINE. Here begins Wolff's articulated disagreements with Haller over whether development occurs gradually and epigenetically or by unfolding of preformed material. "I consider it proven that the intestine is doubtlessly thus formed (by rolling of material) and did not exist previously in an invisible form, ready to appear at the appropriate moment, "wrote Wolff in opposition to Haller. Just because he could not see the parts early on does not mean that they could not exist already, Wolff realized, but he believed that in fact the parts are only formed as the result of a gradual process. Unfortunately, Wolff's work was little known, even after a translation into German appeared. Only much later was Wolff appreciated, according to Blyakher, and it took figures such as von Baer, the American biologist William Morton Wheeler, and the embryologist-historian of science Joseph Needham to evaluate properly Wolff's "fatal blow for preformation."


CHAPTER 6 considers Wolff's teratological work, performed after his move to Russia and based on the St. Petersburg collections in the Kunstkamera. These studies, published in Latin, have remained essentially unknown until recently. After arguing that God would not have created monsters, Wolff maintained that abnormalities must occur by epigenesis rather than preformation. Blyakher asserts that Wolff's discussion of God reflected his desire to "eliminate God from nature" and that any impression otherwise stems from Wolff's necessary conformity to prevailing popular opinion. But the reader should consider this claim sceptically, a warning reinforced by awareness of Blyakher 's efforts through the last few pages of the chapter to make Wolff a predecessor of modern embryology.

CHAPTER 7 presents Wolff's "essential power" as discussed in his commentary for the 1782 St. Petersburg Academy of Science prize competition for understanding nutritional power. In a paper of his own, Wolff responded to papers by Blumenbach and Born by discussing attractive and repulsive forces and the importance of forces as well as structure for organic animal development.

CHAPTER 8 addresses evaluation of Wolff's work by Kirchhoff (Wolff brought development from mystery to a science by establishing that organic life follows laws) and Raikov (Wolff was a materialist and denied the existence of Stahl's mystical "soul," an idealist but not a vitalist, stressing the primacy of material over soul) . Interestingly, given his retrospective tendencies elsewhere, Blyakher believes that Raikov distorts the proper historical perspective, and he sees Wolff as fluctuating between materialism and idealism. Consistently, Blyakher tries to show how major figures were predecessors of modern science even though they were sidetracked by errors of their day. Thus he is Whiggish in his history, but he is not completely ahistorical. It is not Wolff's fault that he could not do more, Blyakher apologizes; the backward times slowed Wolff's progress in Blyakher 's assessment. Thus, like earlier chapters, this chapter begins with useful description and references to relatively littleknown material and ends with a claim for Russian priority.


CHAPTER 9 argues that Wolff was essentially ignored but that Russian embryologists nonetheless began to accept epigenesis by the late eighteenth century. Blyakher discusses such figures as Johannes Beseke, Matvei Pekken, Nestor Maksimovich-Ambodik, and Aleksandr Radishchev, providing a valuable, though brief, introduction to each of these scientists.

CHAPTER 10 is perhaps the most significant in introducing a cast unfamiliar characters and unfamiliar material, and in providing original theoretical discussion. After establishing what Naturphilosophie means to him, Blyakher assesses the impact of German Naturphilosophie on Russian science; he concludes that Russians were generally not receptive to Schel ling's philosophy or to idealistic Naturphilosophie in general, even though some embryologists such as Danil Vallanski, Michael Pavlov, and others endorsed seemingly idealistic views. The Russian intelligentsia recognized the unreality of Naturphilosophie and the importance of materialism, Blyakher argues, and thus they moved toward a progressive empirical philosophy. Despite apparent flirtations with Naturphilosophie, therefore (as for von Baer) , Blyakher concludes that "the successful aspects of embryology in Russia were thus not connected with Naturphilosophie." Although once again consistently retrospective and apologetic for the seemingly imperfect progress of Russian science, Blyakher has in this chapter addressed the suggestion by others that Naturphilosophie may have directed Russian science and argues that it may have been seriously considered but then rejected or refined in "successful" Russian science. His discussion of those who did accept some form of idealistic philosophy is useful, as is his interpretative assessment of its limits.

CHAPTERS 11 AND 12 sketch, respectively, the contributions of transition figures Louis Tredern and Ludwig Bojanus. Tredern admittedly "was not a Russian, was not born in Russia, and lived there only six years." Yet he was an honorary Russian in Blyakher' s view. Tredern did produce an influential dissertation, reportedly inspired by the Russian Wolff and by Tredern 's visit to St. Petersburg; there he outlined the preliminary story of the avian egg and its hatching and early development. Bojanus introduced study of the embryonic layers in mammals, which influenced Pander and von Baer, according to Blyakher.


CHAPTER 13 discusses Khristian Pander, von Baer's fellow student at Wurzburg studying under Dollinger. Dollinger and von Baer convinced Pander to apply his apparently significant financial resources to procure the necessary large number of eggs in order to trace details of chick development during the first five days of life. Pander's work, despite criticism by Lorenz Oken which Blyakher discusses in detail, provided a starting point for future study in epigenetic developmental biology, and especially notably, it served as a foundation for von Baer's work. At one point, Blyakher almost perversely manages to make Pander's weaknesses sound like strengths. He says that Pander's errors were valuable and that they were important in part because they later "allowed Baer to give the true interpretation." As before, Blyakher 's interpretation remains retrospective and frustrating at times, but his data are useful for an introduction to this material.

CHAPTER 14 THROUGH 24 deal with Karl Ernst von Baer, here Karl Maksimovich Baer. 14 provides biographical information and outlines his professional career. 15 presents Baer's discovery of the mammalian ovum and reveals concern both with establishing Baer's priority and with opposition to Baer's work. CHAPTERS 16 THROUGH 2 2 describe Baer's opus, UBER ENTWICKELUNGSGESCHICHTE. Originally published in Germany (volume 1 1828, volume 2 1837, volume 2 part 2 1888), Baer's work appeared in Russian translation only in 1950 and 1953, which may have provided one stimulus to Blyakher to publish his historical study. Blyakher evidently relied on the Russian translation, so I have had to provide references to the German original (as mentioned above). Few people have read through Baer's long and detailed study completely, so Blyakher 's discussion of all five scholia and corollaries and of the rest of the work, of which many are aware but which few read, will prove useful.

Most important, though, is the discussion of Baer's volume 2, and especially in CHAPTER 2 2 of the fourth part which forms the second part of volume 2. This section was published not by Baer himself but by Ludwig Stieda, after Baer's death. Baer had not completed the work, and Stieda discovered the manuscript while working through Baer's materials in order to produce a biography. Baer had begun his study of human development, discussed in this part four, in Konigsberg in 1834, but his move to St. Petersburg that year disrupted his work and he never completed his examination of human normal and abnormal development.

Human development also forms the subject of part of CHAPTER 23, which deals with Baer's teratological work in St. Petersburg. Here Blyakher addresses Baer's complaints about "lack of consideration or unfair attacks, with which his remarkable discoveries were met in Prussia." The Russians were more sympathetic, of course, according to Blyakher. In part because the Germans did not fully acknowledge the importance of his work, Blyakher establishes convincingly, Baer returned to Russia and gave up his systematic embryological studies, turning instead to anthropology and other scientific and family ventures.

The few studies of fertilization and embryological development which Baer did perform after his move, Blyakher discusses, including several papers detailing what is essentially meiosis and mitosis, according to Blyakher. If fertilization and cell-division initiate development, then there could be no pre-existence of individuals; the unfertilized ovum must contain latent but not pre-formed life, Baer had concluded in a paper of 1847. Some of Baer's teratological and fertilization studies reveal that Baer accepted a limited version of evolution - an evolution of the individual within his system of types. Blyakher neatly illustrates the transition between his second and third historical periods of embryology with the example of St. Petersburg Academy of Science's establishment of a prize for Biological Science in 1864. Kovalevsky and Mechnikov won that prize, thus bridging the move from Baer's older epigenetic work to the new evolutionary embryological science.

CHAPTER 2 4 considers Baer's theoretical views, including a very brief look at Baer's version of the history of science. This chapter offers intriguing suggestions, but most are incompletely developed and hence do not significantly extend our understanding of Baer. As with the rest of the book, the chief value of these lengthy chapters on Baer lies in the potential of their suggestions, in the descriptions of more well-known sources and of unfamiliar material alike against a background of other familiar works. The references provided certainly suggest that Baer is as yet poorly understood, despite the several biographical sketches, and that historians of science would do well to explore his complex Russian connections - both before his move to and after his return from Germany.

CHAPTER 2 5 serves as a transition to the third stage of world and hence Russian embryology (featured in Blyakher's second volume) . It considers figures after Baer but before Mechnikov and Kovalevsky. The focus is on Grube, Nordmann, Warnek, and Krohn in particular. These men made way for Kovalevsky and Mechnikov, according to Blyakher, and these latter men effected the revolution from comparative-descriptive to comparative-evolutionary embryological science.


NOTES - These notes have not been translated, obviously. Some offer biographical information, others provide references to additional scientific and other works, while still others elaborate on the text. These notes are cited in the text by the numbers enclosed in square brackets: (#) .

The above brief outline sketches Blyakher's volume. Throughout, the work remains descriptive. Each chapter thus provides details of the works and people it considers. Some of these descriptions are so extremely thorough as almost to reproduce the original sources being considered, while others provide essentially an index or overview of their subjects. To my knowledge, the descriptions seem consistently reliable and useful.

Value of the Work

As suggested above, Blyakher's work contributes both useful description of little-read source materials and a particularly Russian perspective. The latter, which clearly directs what interpretation Blyakher does offer, only occasionally intrudes on the narrative. As noted earlier, Blyakher does at rare times become fervent in his attempts to establish that "Russia is the fatherland of embryology as a science." Yet he would appear to have considerable evidence that his claim should at least be taken seriously. Western scholars often tend to dismiss Soviet scholarship and its fiercely patriotic perversions. But Blyakher, despite his effectively cold war context, remains relatively restrained and reasonable.


The author's concern with establishing scientific priorities, with establishing who first discovered such-andsuch, seems equally open to objection from the perspective of current history of science. Yet this orientation clearly does not result strictly from Blyakher ' s Russian point of view; most historians of science in the 1950' s sought to establish priorities and to document high points of scientific "progress."

In sum, then, Blyakher does provide a very useful descriptive guide to major works in the history of embryology, many of which happen to be Russian in some sense. His interpretative discussion, which seeks to establish that the Russian connection in important embryological work was not merely coincidental needs to be questioned, dissected, and then explored further to discover just what the essential Russian influences were. We should thank Blyakher for his suggestions and use his volume as a guide for that further exploration.

Because the materials are so widely known, I have decided not to provide a full bibliography of works relevant to the subjects Blyakher discusses. See the DICTIONARY OF SCIENTIFIC BIOGRAPHY entries for the key figures and standard sources in the history of developmental biology for additional references and for discussion of similar materials from various non-Russian perspectives.

Jane Maienschein

July 1981 Arizona State University


Preface

Embryology occupies a notable position among the biological disciplines of Russian science, as noted by K. A. Timiriazev. 1 Separate stages of the history of Russian embryology are presented in journal articles, collected biographical essays, and commentaries which have accompanied the recently published works of the Russian scientists K. F. Wolff, K. M. Baer, A. 0. Kovalevsky, and I. I. Mechnikov There is no systematic treatise of the history of embryology, either here or abroad. 2 Joseph Needham's HISTORY OF EMBRYOLOGY, which was translated in 1947 into Russian, 3 cannot satisfy any exacting reader. (1)4 For exhaustive evidence of the frequently repeated claim that Russia is the fatherland of embryology as a science, that it developed from Russian soil and became one of the most important foundations of the evolutionary and historical view in the organic world, therefore, it is necessary to give a detailed account of the history of Russian embryological investigations K(liment) A(rkadevich) Timiriazev, "Istoricheskii metod v biologii" (Historical methods in biology) .... SOCH., V.VI.M., Selkhozgiz (1939), p. 32.

This confirmation is correct now only in relation to the history of embryology of animals . After the manuscript of this present book was given to the press , there appeared P(avel) A(leksandrovich) Baranov's monograph, ISTORIYA EMBRIOLOGII RASTENII V SVYAZI S RAZVITIEM PREDSTAVLENII ZAROZHDENII 0RGANIZM0V (History of plant embryology in connection with developmental ideas on the generation of organisms) (Moscow: Akademii Nauk, 1955, 439 pp.) . In the first part of this book its author presented the prehistory of the embryology of plants , to the nineteenth century, against the background of development of general ideas on the generation of organisms, both plant and animal (p. 8) J. Needham, HISTORY OF EMBRYOLOGY, translated from the English by A. V. Yudinaya, preface by V. P. Karpov (IL, 1947) , 342 pp.

Numbers in square brackets are related to the comments at the end of the book .


The first observations of embryonic development date back 2,500 years; however, the onset and development of embryology as a science is connected with the general progress of the natural sciences in the eighteenth and nineteenth centuries. There are three main stages in the history of embryology. The period of the substantiation of the theory of epigenesis, making possible the existence of embryology, is connected with the Russian academician K. F. Wolff. The period of the discovery of the embryonic layers, when the prerequisites for comparative embryology appeared, is the period of activity of the Russian academicians Kh. I. Pander and K. M. Baer. Finally, the period of the creative development of embryological problems in the light of Darwin's work is inseparably united with the Russian investigators A. 0. Kovalevsky and I. I. Mechnikov and the brilliant zoologists who followed their footsteps.

The present book reviews the first two periods of the history of embryology, including the epoch directly preceding the scientific activity of A. 0. Kovalevsky and I. I. Mechnikov. The history of Russian embryological investigations, from the beginning of comparative embryology to the present time, is embodied in the contents of the books discussed here. One is dedicated to the history of embryology of invertebrates and another book elucidates the history of the embryology of vertebrates.

At the time of writing the present book, no work of Wolff or Baer had been published in Russian except for the incomplete and inaccurate translation of the second part of the first volume of Baer's UBER ENTWICKLUNGSGESCHICHTE .5 In 1950, in the academic series "Klassiki nauki," the translation of Wolff's "Theory of Generation" appeared, as did the translation of the two volumes of Baer's "History of Animal Development" in 1950 and 1953. An article by Gaissinovich 6 (in an appendix to Wolff's book) clearly presents this embryologist 's ideas against the historical background of studies of evolution. An analogous article by B. E. Raikov concluded the first volume of Baer's work. 7 The appearance of these articles has reduced the corresponding chapters of the present book, but the author considers it necessary to give a brief, thorough review of the basic works of Wolff and Baer in connection with their other embryological and teratological works.


5. K. M. Baer, IZBRANNYE RABOTY (Selected works), from the series KLASSIKI ESTESTVOZNANIYA, translated with comments and preface by Yuri A(leksandrovich) Filipchenko (Moscow: GIZ, 1924), 114 pp.

6. A. E. Gaissinovich, "K. F. Wolff and Studies on Development," in an appendix to K . F. WOLFF: THEORY OF DEVELOPMENT (Moscow: Akademii nauk, 1950), pp. 363 - 477


The works of Russian embryo logists of the eighteenth and the beginning of the nineteenth centuries — Wolff, Tredern, Pander, M. G. Pavlov, and Baer — are, in part, written in Latin and cannot be read in the original by the majority of our young contemporaries. The literary legacy of Baer is extremely vast and difficult to review.

It seemed to the author that it is important to explain not only the most important works of the Russian embryologists of the second half of the eighteenth century and the first half of the nineteenth century, but also their less important works, in order to present the contribution which Russian investigators made to world science. The activity of the Russian embryologists coincided closely with the development of world embryological science. The importance of the Russian investigations can be correctly evaluated only in comparison with the works of the foreign embryologists, which are explained by short comments in the present book. Expansion of these excursions into the history of world science did not seem possible without significantly increasing the size of the book.

In presenting the contents of the Russian embryological works, sometimes sufficiently long extracts are presented that the reader can form an impression not only about the contents, but also about the scientific literary style of the investigations discussed.**


7. B(oris) ECvgen'evich] Raikov, "On the Life and Scientific Activity of K. M. Baer," in an appendix to K . M. BAER: THE HISTORY OF ANIMAL DEVELOPMENT, Vol. I (Moscow: Akademii nauk , 1950), pp. 383 - 438.

8. All the citations printed are underlined by the authors of the discussed works permitting the reader to see exactly what the authors, themselves considered most essential in their works .


The drawings are considered to be the most important part of the morphological works. The pictures, reproducing the studied objects as they were seen either with the naked eye or with the aided eye, reveal the level of the technique and the exactness of the observations. The schematic drawings are considered the graphic expressions of the authors' theoretical opinions. Therefore the original drawings speak about the scientific priorities and facilitate the description in words of the corresponding discoveries. In reference to this, we must not forget the authors of revisions and educational textbooks; although they occasionally reprint the illustrations of the foreign authors, often they are not a bit better than the drawings of the Russian investigators published earlier. The reprinting of the original drawings of Wolff, Tredern, Boyanus, Pander, Baer, Grube, Nordman, Warnek and Krohn in the present book is to present the level of their morphological investigations, and also to defend the priority of the Russian embryologists .

The present book was finished in 1950. Its contents, and also the plan for its continuation to subsequent stages of embryology in our country, were reported in the meeting of the sector of the history of biology in the Institute of Natural Science History of the Academy of Science of USSR and in the Academic Council of this Institute.

The author is deeply grateful to T. D. Detlaf; L. D. Lioznera, S. R. Mikulinsky, S. L. Sobol and G. A. Shmidt, who listened to the reports about the contents of this book or read the manuscript and made critical comments. The author particularly thanks the collective of the library of the Moscow Society of Naturalists, who willingly helped during the search for literary sources and illustrated materials.


Comments

(1) Needham arbitrarily limited the contents of his book absolutely to the history of chemical embryology. In addition, his extreme unobjectivity drew attention, as he dwelt mainly on the works of English authors, while, according to his opinion, the book should have represented the history of embryology of all times and peoples. In the chronological table of scientists of the world embryology, covering up to the beginning of the 20th century (p. 266 of Russian translation) , Needham did not find places for the names of N. A. Warnek, A. 0. Kovalevsky, I. I. Mechnikov and their numerous Russian followers in contrast to

Fr. Balfur, who published more than 70 years ago the first (and for that time excellent) manual book on comparative embryology, in which the Russian works were given proper places (14) .

(2) Samples of these "compositions" are pictured on the prints of that time, reprinted in the book "Kunstkamera Peterbergskoi Akademii Nauk", 1853, 293 p. (see Fig. 6.10 and 11 of the mentioned book) . Description of anatomical and embryological "compositions of Ryuish and texts of the poetical Latin inscriptions which accompanied them much earlier were mentioned by Baer in "Memories about anatomical cabinet" ("Memoire uber das Anatomische Kabinet, gelesen in den Sitzungen der Phys.-math. Kl. d, Sep. 20, 4 u. Oct. 18, 1850; Collection the museum of anthropology and ethnography in Imp. Ac. Sc, 1900, p. 111-152) (21).


Here and below figures in round brackets show the number of text page, to which the comment is related.


(3) Doctor van der Hulst, (see below for data about him) on April 14, 1724 ,; sent organs which belonged to a girl, who died of smallpox, in Kunstkamera. Caesar honored him due to the hermaphrodite (Materials for history of Imp. Ac. Sc, V. 1 Spb, 1855, p. 38) in the other place of these "Materials" wrote due to bringing to the Academy of Science a monster, who was born with two heads, determined by Doctor Stepan Gaokantsk, ...8 roubles were given" (V. II, on March 9, 1732, p. 119) (23).

(4) In "Materials for history of Imp. Ac. Sc." there are many documents about the obtaining of teratological objects into Kunstkamera.

A letter from Vyborkh by colonel general Karnov says "his major Beshentsev Fedor Fedrov in the Vyborgskaya office found a monster of a lamb having four legs and one neck, and the head had the view of a double one. This head had two mouths with tongues and two eyes: they were in the

middle where the forehead must have been " ('Materials..

V. I, Spb., 1855 No. 165, March 9, 1725, pp. 9-97).

Inventory of objects, obtained on March 8, 1725 "Number 1: lamb, with 8 legs, another with three eyes, two trunks, 6 limbs were sent from Tobol'sk by Kozlovsky. Number 2: baby, with 3 legs: from Lower Novgorod from Governor Rzhevsky. Number 3: calf with 2 deformed legs: from Ufa from commandant Bakhmetov. Number 4: baby with two heads also from Bakhmetov. Number 5: one baby, with eyes under the nose and ears under neck: from Nezhin. Number 6 - two babies - breasts and abdomens were joined from Akhtyrok from prince Mikhail Golitsyn; hands, legs and head were normal. Number 7 - baby, with a fish tail, born in Moscow in Tverskaya. Number 8: two puppies, born from a 60-year-old woman, from Akhtyrok from prince Golitsyn. Number 9: baby with 2 heads, 4 hands, 3 legs: from Ufa from commandant Bakhmetov" ("Materials...", V. I, No. 193, March 18, 1625, p. 99).

Grigorii Ivanov found a monstrous head puppy with 8 legs ("Materials, V. I, No. 291, September 21, 1725, p. 145).


"....Dead body of an infant, with 6 fingers, horn by a daughter of Tikhanov who was working on a marine ship Astrakhan. . .given in Kunstkamera" ("Materials..." V. II, 1886, No. 99, November 29, 1731, p. 83).

"On January 17 of this same year, my brother sent a monster which had 2 mouths, the daughter of Ulita Kiryanova the wife of sergeant Nikivor Kosharov gave birth to it: this monster had four hands, four legs and two faces, one had normal view and the other had one eye" ("Materials...", V. Ill, 1886, No. 13, January 20, 1736, p. 14).

"This monster was accepted for preservation in Kunstkamera and Kosharov was given 4 roubles as a reward" (there is also, No. 41, February 17, p. 36) (23).

(5) Iogani-Georg Dyuvernua (1691-1759) a Russian academician in the department of zoology and anatomy, was invited to the academy during its foundation. In 1741, by efforts of Shumakher, he was compelled to leave work and travel to his homeland, in Germany. Following him, up to 1758 Abraham Kaau-Boerhave (1715-1758) had been in charge the department of anatomy and physiology in the Academy of Science. In his era, the anatomical and embryological collections of the academy came into being, but he practically did not use them. He published one of them in detail (Abraham Kaau-Boerhave was related to those foreigners, living in Russia, who did not make anything for promoting its sciences and for preparing the young Russian scientists (23) .

(6) Family of van der Hulst was repeatedly mentioned in the documents of Petrovskaya epoch. Doctor Zakharii van der Hulst arrived in Moscow from Holland, apparently in the 70s of the 17th century, and for a long time he was the physician of Aptekarsky department and court physician at tsars Ivan and Petr. Then after the death of Ivan, he became the physician of Petr I. He accompanied him on both journeys to Arkhangelsk (1693 and 1694). On the second Journey, he die-' suddenly* . Another van der Hulst participated in 1691 in Poteshny battles" in the army "generalissimusa" of I. I. Buturlin 2 . In 1695 in "Rospisi nachalnym lyudam Semenovskogo polky" captain Andrei Yakovlevich van der Hulst was mentioned^. On the first journey of Petr to Holland, at the Russian Embassy, lieutenant (or captain) Andrei (Yakovlevich) van der Hulst 4 became the translator. One year later, t he was sent by the Dutch government to Moscow as resident* Later on, the son of the above mentioned "doctor" Zakharii van der Hulst— Zakhar Zakharovich— was known. By the order of Peter I, he received "traveller sheet" via Mozhaisk, Vyazm, Dorogobuzh and Smolensk in foreign lands for studying science. Although it was impossible to detect exactly the time of his return to Russia, but there was no doubt about it . He was a teacher of surgery at Peterburgsky Hospital 1723 6 . Ya. Chistovich informed, that "Z. Z. van der Hulst passed the Doctorate degree examination in Leiden and after returning to Russia he was the senior doctor in Petersburg Admiral Hospital and, in addition, a teacher for medical students and pupils of this hospital. Later on, he lived in Moscow and when a "Doctor's committee"? (1730) which comprised five doctors was established, he was one of its members. It is most probable, that Zakhar Zakharovich van der Hulst is the author of a dissertation cited in the text on page 7: the difference of names (Zakharii and Arnold) does not speak against this supposition, as both names could belong to the same person. Anyhow there is no doubt about the belonging of the author of dissertation to the Moscow family of van der Hulst (25) .


V. Rikhter. "Istoriya meditsiny v Rossii" (History of Medicine in Russia), p. 2, Moscow, 1820, p. 313, M. M. Bogoslovskii, Petr I, V. I, p. 181= V. II, 1941, pp. 123-124.

2. There also, V. I, p. 127.

3. "Sb. vypisok iz arkhivnykh bumag petre Velikom" (Collection from archives papers about Petr the Great) , VI, Moscow, 1872, p. 148.

4. M. M. Bogoslovskii. Petr I, V. II, p. 155, 182, 421.

5. There also, V. II, p. 432> V: IV, 1948, p. 252, 339, 340, 344, 346.. M. A. Venevitinov. "Russkie v Gollandii. Velikoe posol'stvo 1691-1690 godov" (Russians in Holland. The great embassy 1697-1698) Moscow, 1897, p. 79.

6. V. Rikhter, Istoriya meditsiny v Rossii, p. 3, 1820, p. 149.

7. Ya. Chistovich. "Istoriya pervykh meditsinskikh shkol v Rossii" (History of first medical schools in Russia) . Appendix X. Alphabetical list of doctors of medicine, working in Russia in the 18th century.


(7) Speaking about the collection of anatomical preparations, present in Kunstkamera N. G. Kurganov noticed, that

"the greatest attention was given to these parts, which explained parturition. A number of the foeti exceeded more than one hundred and composed a gradualness from an embryo having the size of an anisic grain to a completely formed baby. The collection of monsters was extremely big. These anatomical descriptions with sketches, reprinted on copper, had a scientific significance" ("Pismovnik", the second part, p. 196) (32) .

(8) In chapter 26, "Gippokratovskoi sbnornik" it is possible to read the following: "All organs are distinguished simultaneously and they grow, and not one is distinguished earlier than the other. But the larger ones in nature are distinguished before the smaller ones, not originating in any case earlier. However not all receive the final structure in equal time, but some are quicker, others are slower, since each meets sufficient nutrition. In some, all become distinct within 40 days in others — within 2 months, in others " within 3 months, and in others " within 4 months" (See V. I. Karnov. Aristotle i antichnaya embriologiya. Introductory article in the translation of Aristotle" "0 vozniknovenii zhivotnykh" Izd. AN SSSR, 1949, p. 23 (35).

(9) The embryological opinions of Dekart are stated in the treatise "Opisanie chelovecheskogo telo" (Description of human body) , where they compose its fourth part — "About the development of the embryo. Parts, formed in semen" and the fifth part "the formation of hard parts". The treatise was published two years before the death of the philosopher, in 1648. In addition, with the birth conception of living substances, "which are produced by semen", Dekart also assumed the possibility of spontaneous conception (without semen and uterus") . *

In case of conception from parents, their semen is mixed and forms cloudy liquid which undergoes a kind of fermentation, what is formed during this heat widens the particles of semen, they "press on other particles subsequently locating them gradually. This is the way of forming the body organs... Heat compels some of the particles of semen to be collected near definite points of the space... Thus the heart begins to be formed"2. The movement of blood from heart makes a way through semen particles, that is why the blood again returns to the heart, and by this way the vascular system is formed. After this, the movement of particles of different kinds leads to the subsequent formation of the organs - vertebrates with spinal cord, brain, paired organs of sensation and so on. The energetic character of these embryological presentations is completely obvious. However, Dekart considered that it is necessary to underline this. "In order to get acquainted with the figure of the already formed animal, it should be understood what it represents at the beginning of its formation and it is necessary to imagine semen, as some mass, from which, the heart is first formed, around it the hollow vein is located on one side while on the other the large artery, united by two tips. The tips of these vessels, to which the openings of the heart are directed, indicate the side, where the head must be present, others also indicate that side, where the lower parts of the body must be present" 3 (36) .

(10) The idea of preformation may be traced back to remote ancient times. Anaksagor taught that "hairs cannot be formed from no hairs and raft from no raft" similar ideas were stated by several authors especially Senekoi, who in "Questions of nature" wrote: "In semen all future parts of the human body are contained. The baby in the uterus of the mother has already roots of beard and hair, which he will carry. In a similar way, in this small mass, are contained all features of the body as well as all those, which will be present in his posterity" (see Dzh. Needham, History of Embryology, page 76) (37) .

1. See S. F. Vasilev. "Evolution ideas in Dekart philosophy (Introductory article in book" Rene Dekart. Kosmogoniya . Two treatises. GTTI. 1934, p. 121) .

2. R. Dekart. Description of human body. In book: Kosmogoniya, p. 286-287.

3. See also pp. 298-299.


(11) The theory of "investment", conformable to plants and animals, was stated by Mal'bransh in the following expressions: "It seems, although it may be a hardlyaccepted idea, that in one embryo a countless number of trees is included, because this embryo does not include only a tree, serving as its seed, but also, great number of seeds, which can include new trees and new seeds of trees, containing, in turn, probably in an incomprehensible little form, other trees and other seeds, are fruit bearing as the first and so on till infinity. All what is mentioned about plants and embryos can be also applied on animals and their embryos, from which they were produced. In the embryo of a bulb of tulip, it is possible to distinguish all tulips. Thus in the embryo of a fresh non-hatching egg as well, it is possible to see a chick, which may be nearly completely formed. In the eggs of frogs, it is easy to recognize frogs, we will also find other animals in their embryo when we become so experienced and skilful, that we can open them (N. Mal'bransh. "seeking out truth" translated by E. B. Smelovaya, V. I, 1903, p. 51-52) (38) .

(12). Speaking of the influence of the mother's impression on the formation of fetus, Mal'bransh, among other examples, mentions the following: "One year did not elapse still from that time, when a woman, looking with great zeal on the picture of Saint Pia at the time of the celebration of his canonization, delivered a child, absolutely like the Saint He had senile face, which was impossible for a child, but did not possess a beard. His hands were put together as cross on the chest, his eyes were directed to the sky. He had a very small forehead, because on the picture, the image of this Saint was raised to the dome of the church, directed to the sky, so that his forehead was nearly unnoticed. On his shoulders, he had something like overturned mitre with some round birth marks, at these places where mitres were decorated with precious stones. In short, this infant extremely resembled the picture, but his form was made by the strong imagination of his mother. All Paris could see him, as I could, because this infant was preserved for a long time in spirits of wine" (N. Mal'bransh. Seeking out truth, V. I, p. 173).

The description of deformity was so expressive and exact, that confidently it was sufficiently possible to characterize it in terms of recent teratology. Probably, the matter was a case of amencephaly or, may be, cerebral hernia of occipital region. It is not wonderful that the baby, resembling Saint Pia, was shown in a jar with spirit directly after bith (38) .

(13) Philosophy, Leibnits spoke, - gave itself much work about the origin of forms, entelechy and soul. Meanwhile, different accurate investigations, performed on plants, insects and animals, led to this conclusion: that the organic bodies of nature never originated from chaos or not, but always from semen, in which, undoubtedly, preformation was already present... We see something similar, when, for example, worms become flies and caterpillars become butterflies" (Monadologiya, No. 74), (38).

(14) "Thus, I suppose that souls which once must become human souls, as well as souls of other kinds, existed in semen, in ancestors up to Adam, i.e. from the very beginning of things they existed in the form of organic bodies — a view, which was apparently approved by Swammerdam, Mal'bransh, Beil', Pitkari, Gartsuker and many other learned men. This view is also sufficiently confirmed by microscopic observations of Leeuwenhoek and other fairly prominent naturalists "(Teoditseya, I, No. 91) (38).

(15) Leibnits considered that monads are alive and animated, characterized by incessant change which is accomplished continuously without leaps. "I confirm, as an indisputable truth — he wrote — that all things were exposed to change, and became monads and that in each monad this change was accomplished continuously" (Monadologiya, No. 10) . From continuous change, the developing monad natural passage of Leibnits to gradation of monads, forming continuous eternal ascending series of substances, progressing from unaccomplished to accomplished. By his investigations "Swammerdam showed that insects, by their respiratory organs, are similar to plants, and that in nature, an order of gradualness, descending from animals to plants, exists. However, there may be, in addition, intermediate substances between these and others" (Letter of Leibnitz to Bung). And in another letter: "I am sure, that these substances must be present, and natural science may discover them. Nature never disturbs continuity anywhere. It does not make leaps. All categories of substances of nature form one sole chain, where different classes, like links, so closely join to each other that for sensual presentation it is impossible to determine the point, where each of them begins or ends" (cited by Kuno Fisher) ("History of new philosophy, V. Lebnitz", p. 460) (38) .

(16) About this change of his opinions, Haller wrote the following: "In the body of animal, there is no part which can originate earlier than the other: all of them are formed at the same time... If Harvey supposed that

he discovered the epigenetical development, it is because from the beginning he only saw small haziness and then the rudiments of the head and eyes, exceeding in size all other bodies, and finally gradually — the internal organs. More than 20 years ago, i.e. before my numerous observations on eggs and females of Tetrapoda, I used this argument to prove that embryo strongly differed from the formed animal, while I confirmed, that in animals at the moment of conception parts which were present in completely formed animal were absent. From this time, I had the complete possibility to confirm that all that was deduced by me against preformation theory, in fact speaks in its favor" (cited by Dzh. Needham, History of Embryology, p. 226-227) (39).

(17) On the basis of the confirmation of the Bible, that earth and mankind populating it, have existed about 6000 years and from that the average duration of life of

man is equal to 30 years, Haller calculated, that God created at the same time a minimum of 2000,000,000,000 people (cited by article of Kirchhoff on Wolff, p. 204) (39) .

(18) Speaking about the impression produced on contemporaries by the discovery of Bfennet (about the development of bodies from unfertilized ovum), A. E. Gaisinovich noticed that the scientists of that time... lost sight of that it proved in the best case only ovism but not preformation (Cited article in the edition of translation "Theory of conception" of Wolff, 1950, p. 379). With the latter confirmation, it is difficult to agree: ovism is one of two forms of the preformation theory and is principally identical with its other form — animalculism. (40)

(19) After that the chapters of this book, dedicated by Wolff, were written, he mentioned in the preface "Theory of conception" of Wolff (Publisher House Ac. Sc. USSR, 1950) which was published with the supplement of article A. E. Gaisinovich "K. F. Wolff and studies on development" (pp. 363-477) Tasks, which the author of this interesting article put before himself, did not allow him, apparently, to stop and dwell in more detail on embryological and teratological works of Wolff.

Here it is also necessary to notice that the translation replaced in the Russian edition of Wolff the term "generation" by the word "conception", which could not be considered felicitous. The term "conception" meant the beginning stage of development, appearance of a new individual, while Wolff did not mean only this stage, but also all subsequent individual development. It is completely accurate to translate "generation" by the word "development". Wolff himself used in his German book neither the term "conception" (Entstehung) nor the term "development" (Entwickelung) , and dept the Latin root in the German word "die Generation". In accordance with this, upon the examination of Wolff's dissertation and its popular summary in German below the original Wolff term "generation" is preserved (43) .

(20) In the first volume of "Zur Morphologie" Gete mentioned Wolff in four places. He gave a brief account of biography of Wolff (p. 80-83), mentioned in the notes of Murzinitsa on Wolff (p. 252-256), then dwelt on studies of Wolff about metamorphosis of plants (p. 83-87) . At last he tackled the understanding of educational yearning, using the terms of Blumenbakh (pp. 114-116).

In the last extract Gete writes the following "In criticism of ability to reach an opinion" Kant states "In relation to the theory of epigensis no one worked either for its proof and for substantiation of true principles of its application, or partly for the restriction of its extremely wide application, as Mr. Buildings advisor Blumenbakh". This evidence of honest Kant encouraged me again to scrutinize work of Blumenbakh, which I truly read before, but did not imbue it. Here I found my Christof (1-L.B.) Friedrich Wolff as an intermediate link between Haller and Bonnet; on the one hand, and Blumenbakh, on the other. For his epigenesis, Wolff must suppose the presence of an organic element, on which creatures feed, which is intended for organic life, and supplied this material by an essential force". The mentioned words of Gete witness to the superficial acquaintance of the great poet with the views of Wolff (Even Wolff's name was written incorrectly by Gete (44) .

(21) A. E. Gaisinovich (1950, p. 462-463) mentioned information, that Wolff at the beginning of the 70th year taught in the academic high school— chemistry, anatomy and botany, and he also directed the preparation of the scientific activity of student Fedor Galchenkov (48) .

(22) This place in the translation of Meckel tendentiously stated: instead of "the Highest Creator" was put "The creating nature". Wolff disputed the preformation of formulated parts; in the German translation, his skepticism is not related to the Creator for the authority is untouched, but to nature. This "liberty of translation" bars the radicality of scientific and philosophical view of Wolff (76) .

(23) Here the phrase made by Wolff, has no connection with other discussions and clearly intended for not blaming atheism: eiusmodi vero materia, talibus, viribus instructa immediate a Deo ex nihilo creata sit (it is also true that material, supplied by these forces, is directly created by God from nothingness) (76) .

(24) A. E. Gaisinovich (1950, p. 455) repeated the mentioned statement of K. M. Baer nearly literally: "This remarkable work of Wolff... did not draw the attention of all the scientific world up to 1812, when Meckel translated it from the Latin language". It is necessary to notice, however, that the work of Wolff was given due attention and it was evaluated as a remarkable work in the book of I. Bezeke, published in 1797 (see p. 114) and in the dissertation L. Tredern (1808) (see Chapter 11) (87) .

(25) Apparently, in the declaration of biological works by the Academy of Science for a prize, Wolff showed initiatives also early. Thus, in 1779 the academy declared a competition for a prize on the question about reproduction of cryptogamous plants. This theme of competitive work

was written in expressions which a great probability impel to consider Wolff the author of this question: "Theoriam generationis et fructificationis plantarum cryptogamicarum Linnaei dare etc...." (gives the theory of development and fruiting of cryptogamous plants, by system of Linnae, plants and so on) .

In 1783, this prize was awarded to a professor in Leipzig, log. Gedvig for the work under the title "Theoria generationis et fructificationis plantarum cryptogamicarum Lennaei, mere propriis observationibus et experimentis superstructa dessertatio, quae praemio ab. Academia imp. Petropol. pro Anno MDCCLXXXIII proposito ornata est" (105).

(26) It is necessary to notice, that the embryological problems have interested Petersburg Academy of Science before the arrival of Wolff. It is possible to judge this, in particular, by the published collection in 1756, which is composed of two works (dissertations) , and sent to the competition announced by the academy. The question, put by the academy, touched upon the possible influence of the impression, felt by the pregnant woman, on the developing fetus. One of the mentioned works belonged to a professor

at Leipzig University C. Ch. Krauze*, and another — to a member of Petersburg Academy of Science, I. G. Rederer. In lively controversy on this theme Byuffon participated. He energetically objected to preformists, who according to their views, the embryo is proved to be similar with the parents under the influence of imagination of the mother.

Caroli Christiani Krauze. Dissertatio de questione ab Academia imp. scientianim petropolitana pro praemis in annum MOCCLVI proposita. Quaenam sit causa proxima mutans, corpus foetus etc... There also, J. D. Roederer Dessertatis Petropoli 1956.


In the controversy Terner also participated. He objected to Blondel, denying the influence of mother imagination on the embryos. Argument of Terner was confirmed by the fact that the blood vessels of the mother directly passes in the vessels of the fetus. Ens subscribed to Terner' s opinion. Apparently, the difference, existing on this question in literature, induced Petersburg Academy of Science to announce competition on works which would comprehensively answer it. The scientific committee of the Academy, examining the competitive works, revealed scientific impartiality, and the Academy published two works, each one of them represented a contradictory opinion. Krauze was a supporter, and Rederer was an opponent of the influence of maternal imagination on the fetus.

Krauze considered that the question raised was very difficult, nearly hopeless to solve. However, he noticed, that many examples were present, when the fetus was changed in a way, that not only simple people and the mother herself, but even sharply sensitive scientific people in medical practice related it, although partially, to the strong emotional shock of the mother. Enumerating the opinions of authors, discussing this question (Sennert, Morisso, Ludvig, Hofman, Abraham Kau-burgav, Takhoni and others), Krauze suggested, that the reader can make for himself the most classical ironical defiance: "Hie Rhodus, hie salta!" (literally: "Here Rodos also jump!") and all the following statements attempt to answer this call. As also the majority of his predecessors, Krauze paid attention mainly to cases of appearance of pigmental birth marks, where their forms and situation compel to suggest the influence of pregnant women. He began with these sharp effects, as terror, fear, anger and so on, showing strong physiological influence expressed first of all in the reaction from the side of vascular and nervous systems. From this Krauze made the conclusion, that "if the brain was strongly alarmed, then small changes took place in the body". Referring to known cases of adult people suddently growing grey under the effect of deep feelings, Krauze suggested that, there were more bases to expect changes of skin color of the fetus under the effect of maternal imagination. If terror or fear can cause small ulcers on lips or erysipelas, then why does it seem incredible to you, that the same phenomena can take place in the body of the fetus, whose structure is so weak and delicate, and its vessels are so numerous and full of juices?"


In order to imagine the possibility of this phenomenon, which is shown by the influence of changes in the organism of the mother on the fetus, it is necessary to prove their close relation. On his side Krauze confirmed, "fetus with the uterus represents a single continuous whole". According to this opinion, this is especially related to the nervous system, so that "stimulation of the nerves of the uterus may and must be passed to all the nervous system of the fetus. Distributing this confirmation on psychics, as well as suggesting, that the fetus was capable of psycological manifestations, Krauze deduced the following conclusion: "In the brain of the fetus exists the same condition, which exists in the brain of the mother". All these views are summarized in the concluding paragraph of the dissertation". What was stated so far may be added to all these examples, which instruct, that the fetus body is changed from the mother, if her soul is strongly shocked. Ideas, originating in the mother's brain are "united with fetus brain"; they are stimulated in it more quickly and energetically, than in the soul of the mother herself, as the pulse of arteries and effect of nervous and generally all responses in babies are quicker than in adults. Under the influence of these ideas, the fetus brain gives effect on its body and namely in this way, which corresponds to perceived ideas. Therefore, the fetus's brain produces in the corresponding parts of his body the same things which he himself undergoes".

Directly after the considered work of Krauze, it is necessary to examine the work of Rederer, which is simply entitled "Dissertatis" and also begins with the formulation of the question raised by the Academy: "What is the direct cause for changing the fetus body?..."

In order to possibly answer this stated question, Rederer considered that it is necessary to study thoroughly, how the mother's body unites with the embryo, taking into consideration, that the only connection between them is carried out through the placenta. The investigation of this connection led him to the following conclusion. The lumens of the blood vessels of the placenta, directed to the uterus, up till this stage, are so narrow. They do not admit turpentine oil or any other liquid to pass. True anastomoses, connecting the vessels of the uterus and vessels of placenta, do not exist; Vasa hypogastrica which is full, by all means of a waxen mass or any other fluid, does not pass its contents to the umbilical vessels. This was established either in human carcass, or in living cows, ewes, bitches and other animals. If the pregnant animal is fed on roots of Rabiae tinctorum, then the fetus bones will not be stained with the red color. The blood of the fetus differs from the mother's blood in that it is more liquid. Finally, the fetus pulse also differs from the mother's rate.

Subsequently, the fetus lives its independent life, moving its own blood by its forces, beating of its heart, by blood circulation and through placenta, without the help of the maternal blood. Therefore, rest, movement, sleeping, awakening and even life and death of the mother and fetus are not obligatory present in harmony. Later on, Rederer categorically denied the existence of a nervous connection between the placenta and the mother. The comparison of the positions of the birth marks as well as their presence or absence in the mother and baby does not give, according to his opinion, basis to suggest the influence through blood or nervous system. Rederer discussed in detail the question about "mind" of the fetus and passed to the conclusion, that this mind cannot adequately respond to the feeling of the mother. This is stated by the examples of strong shocks in the mother, not accompanied by the appearance of birth marks in the baby, and examples of appearance of birth marks and warts without any connection with the mother's feeling. In the following pages the various deformities of human beings are mentioned, and Rederer reached the conclusion that the birth marks differ from true deformities only quantitatively. They are also disturbances of the normal development. As the deformities frequently affect the internal organs, then in relation to them, the usual supposition about maternal influence loses its significance. The general conclusion of Rederer comes to that the confirmations about the influence of the mother's feeling on the baby are not supported by verified facts, and are but the product of fantasy (105) .

(27) The effect of vitalistic views of Blumenbackh on some Russian physicians-biologists can be traced up till the 20 's of the 19th century. Thus, in 1825 at Moscow University, the surgeon Nikifor Dmitrievich Lebedev discussed a thesis for the degree of Doctor of Medicine. He later on read at the university "history and literature of medicine". Lebedev dissertation was entitled "About the nature of weightless substances in general and vital powers in particular" (Dissertatio inauguralis physislogica de natura imponderab ilium in genere et de viribus vitalibus in specie, quam...in Universitate caesarea Mosquensi, pro gradu doctoris medicinae. . .elaboravit publiceque defendet chirurgus I-mae classis Nicephorus Lebedev, Mosquae, 1825. 28 p.). Lebedev, referring to Blumenbakh, dwelt on the idea that all vital processes — organic formation and growth as well as movement of already formed parts-possess their own source of a special vital power (55) . The vital power, according to his opinion, is an internal, inherent character in the organic body, which is the cause of life and at the same time seems to be its product (Vis vitalis est interna et proporia organici corpori qualitis, quae vitae causam constituit et simul ejusdem est quasi productum, thesis 9) . Works of similar kind are not characteristic of the common materialistic trend of Russian biological and medical sciences. Therefore, the dissertation of Lebedev served as a reference and mainly as an illustration of this negative influence, which was shown by the idealistic German philosophy and some representatives of Russian sciences especially the naturalists who worked under its direct influence (106) .

(28) Izef Gotlib Kelreiter (1733-1806), botanist and zoologist, worked from 1756 to 1761 as a junior scientific assistant at Petersburg Academy of Sciences, with which he kept in close contact after returning back to Germany and till the end of his life. In the period, from 1758 to 1811, 15 botanical and more than 20 zoological works of Kelreiter were published in Russian scientific and scientific popular editions (Novi Commentarii Acad. Scient. petropolitanae, Acta Acad. Scient. petropol. Nova Acta Ac. Sc. Petropl., Trudy Volnogo Ekonomicheskogo Obshchestva and in the journal "Sochineniya,K poize i uveceleniyu sluzhashchie") . His scientific fame is connected mainly with the study of reproduction and hybridization in plants (see Ioz. Kelreiter" Study about sex and hybridization in the plant", the editor with a biological essay was prof. E. V. Wolff, 1840).


The work of Kelreiter about the irritability in the plant, to which Wolff referred, was called: "Nouvelles observations et experiences sur l'irritabilite des etamines de l'epine vinette (Berberis vulgaris)", Traduit de l'allemand par M. I 1 adjoint Sewergin, Nova Acta Acad. Scient. Petropol., 4, 1790 (German original was received in Petersburg in 1788) (112).

(29) The point of view of A. E. Gaisinovich is similar to the presentation mentioned here about the outlook of Wolff (see Wolff "Theory of conception") . It is possible to combine with it the confirmation, that the studies on preformation and epigenesis do not always correspond to the demarcation between idealism and materialism in biology. The following serves as an evidence, that preformists were also idealists (Leibnitz, Haller, Bonnet) and mechanical materialists (for example, Lamettri) ; equally as epigenetics, were either idealists (Aristotle Harvei) ,

or mechanical materialists (Dekart, Maupertuis, Byuffon and Didar3t).A. E. Gaisinovich included Wolff as well in the latter category. His "absolute epigenesis" was grouped together with Gaisinovich in the mechanical materialism. For its complete verification, this statement could be desired as only a less categorical expression. In the evaluation of Wolff's outlook, it is impossible, apparently, to deny absolutely his fluctuations between materialism and idealism; these fluctuations were an unavoidable originality even by the most prominent thinkers of all historical epochs, preceding the formation of successive system, i.e. dialectic system and materialism (119) .

(30) Semen Gerasimovich Zybelin (1735-1802), after finishing the study in Moscow Ecclesiastical Academy, joined Moscow University till the opening there of a medical faculty. After he finished the course, he was sent to study medicine abroad. At Leiden, Zybelin defended a doctor's dissertation in 1764. After returning back to Moscow he read courses of anatomy, physiology, chemistry, pharmacology and therapy as a professor in the mediqal faculty. In 1784, he was elected member of the Academy of Science (121) .

(31) Prince Dmitrii Alekseevich Golitsyn (1731-1803), a prominent Russian diplomat, was a former ambassador in Holland and France, friend of Diderot and Galvetsy, known as author of many physical works (122) .

(32) Dzh. Needham in his "History of Embryology" prefaced the list of the literature sources used by him with a list of works, which he could not obtain. As "less important works on the history of embryology" Needham also mentioned Bezeke's book. It is difficult to decide, on which basis the English historian of embryology considered Bezeke's book less important, if we take into consideration that according to his confession, he never saw it (122) .

(33) Two extracts compose the contents of the already mentioned book of 1797 (in it, as stated, were present an essay on the history of a hypothesis about the conception and development of animals and in addition, "History of the origin of division of the natural bodies into three kingdoms") The third extract appeared in the form of a separate small volume in the year of the author's death (J. M. G. Bezeke. Allgemeine Geschichte der Naturgeschichte in dem Zeitraume von Erschaffung der Welt bis auf das Jahr N. C. G. 1791. Mitau, 1802, XXXII±154S) . For more details about the works of Bezeke see the article by the author of the present book in "Trudy instituta istoriii estestvoznaniya i tekhniki, V.IV, 1955" (Works of the Institute of History of Natural Sciences and Technique) (123).

(34) Matvei Khristianovich Peken was born in Petersburg, he studied medicine in Ien, where he obtained the degree of Doctor of Medicine (according to other data-- in Gettingen) . When he returned back to Russia he worked as the admiralty doctor, read a course of obstetrics in Petersburg hospitals. From 1793, he travelled to Moscow, where he read pathology and organized the first therapeutic clinic with ten beds (124)

(35) Nestor Maksimovich Maksimovich-Ambodic (1744-1812) finished Kiev Ecclesiastical Academy, studied medicine in Strasburg, where he defended a doctor dissertation about "human liver" (1755). At Petersburg admiralty hospital, he read obstetrics and wrote a lot of works and manual books. The second part of the family (Ambodic) was written by himself in connection with accord of patronymic and family (127) .

(36) "Dictionary" of Maksimovich Ambodic is composed of two parts: Russian-Latin-French and Latin- Russian, which was prefaced with 65 pages of the text, explaining the significance of this first Russian terminological dictionary, contents of anatomy and physiology and even some information about these sciences.

In "the foreward to the dictionary in general", Maksimovich-Ambodic writes: "During collection of the words, related to my subject of practice, I have been collecting bees for more than 10 years. The major reason was that the Russian words had been collected from various ancient and recent manuscripts belonging to church and civics."

It is difficult to overestimate the significance of this work as a result of the great quantity of terms created, which were absent in the Russian language. This significance is not diminished by the bulkness of some terms which are not contained in scientific language together with others which are archaic and have disappeared from the language in the process of its evolution.

The historian of national medicine Ya. A. Chistovich drew a special attention to the significance of "the dictionary" of Maksimovich-Ambodic. He noted the unfair relation to Maksimovich on the part of V. M. Richter, who "did not offer him a single line in his biographical History of Medicine in Russia". Fortunately, the voice of this light word was fair and, in defiance of the partial historian, preserved the name of Maksimovich from undeserved oblivion" (Ya. Chistovich. First obstetrician schools in Russia (1754-1785. Essays from history of Russian medical institutions of the 18th century. SPb., 1870, p. 199) (131).

(37) Khristian Elias Genrikh Knakshtedt was born in Braunshveig in 1749, studied surgery in Bryunn and in 1786 travelled to Petersburg, where he was professor of anatomy and surgery at Kalinkinsky hospital. In 1790, due to the work "Beschreibung der trockenen Knochen des menschlichen Korpers" (SPb., 1791) the medical college awarded him the degree of Doctor of Medicine. Knakshtedt died in 1799. In addition to the mentioned works, he also published "Descriptio praeparatorum maximam partem osteologicorum rarissimorum"


(Braunschweig, 1785) and Latin-German terminological medical dictionary "Erklarung lateinischen Worter, welche zur Geliederungslehre Physiologie; Wundarzneywissenschaft und... Ordnung" (Braunschw., 1784, 2nd edition, SPb., 1788). In the title page of the latter he called himself Russian surgeon and ordinary teacher of studies about bones and all their diseases at the imperial surgical school in Petersburg.

To the work of Knakshtedt mentioned in the text "Anatomical description of the monster" an invitation to Medico-surgical school is addressed to "all famous persons and members of medico- surgical sciences" to attend the meeting "near Kalinkinsky bridge at Ekateringofskaya and listen to "some works and speeches by some teachers and students. The meeting was proposed to take place on January 7 at 10 a.m. (131) .

(38) Petr Andr^evich Zagorskii (1761-1845) is the one who founded the first Russian anatomical school . After teaching at the Cheringovsky college he went to the hospital school in Petersburg. After he finished there, he worked for three years as prosecutor Petersburg medico-surgical school with professor N. P. Karpinsky. From 1799 to 1833, he was junior assistant, and then professor in medico-surgical school, and from 1805 up to the end of his life he was a member of the Academy of Sciences. Zagorskii organized excellent anatomical museums at the medico-surgical school and Academy of Sciences, published the first Russian manual book on anatomy and a great number of works on anatomy, teratology and different medical topics.

(38a) The same opinion on the origin of monsters was also supported by the academician N. Ya. Ozeretskovskii, who informed the Academy on April 25, 17991 about two cases of double monsters in the preparations of the academic museum of natural history. One of these cases was united twins (union in the region of the upper part of the chest, both partners were completely formulated) . The second case was the doubling of the head end, beginning from the girdle region; the monster had three correctly- formulated hands, two normal and one underdeveloped legs. The description of the monsters (the first case was illustrated by two excellently-engraved figures, representing the twins from the front and from behind) was concluded with a brief account about the reason for the monsters appearance. By comparison of the described cases, Ozeretskovskii made the conclusion that monsters can be very variable and that each monster, must possess its natural cause. "The physiologists — Ozeretskovskii wrote — must explain these causes and find the specific original source of these monsters — whether their origin is due to the union of two embryos, or from strengthened, weakened or incomplete development of parts of the body" (p. 371).


1. N. Ozeretskovskii. De doubus foetibus humanis, monstrosis. Nova Acta Acad. Sc. imp. Petropol., 14. 1805, p. 367-372. The article of Ozeretskovskii was published in the same volume of Nova Acta Academiae Petropolitanae , where the above mentioned report of Zagorskii was also published.


In another article, whose contents were received by the Academy one year before (on October 25, 1798)1, a description was given for non-hatched hen's egg with an opening in the shell; through this opening the end of a blood vessel passed. When the egg was opened, in addition to yolk and egg white, a pear-shaped body was found in it, which was full of clotted blood. Ozeretskovskii considered the described content in the egg as a polyp of the oviduct, torn at the time of yolk passing, falling in the egg together with the white. In connection with this, he assumed, that the presence of similar kinds of strange bodies in the eggs of birds may be the cause of appearance of monsters, as the mechanical pressure on the delicate parts of the developed embryo inevitably leads to their deformation. During this, Ozeretskovskii referred to cases of development in double-yolk eggs of doubled embryos which — as a rule — are monsters. During more or less normal development of these twins, they can unite with each other. "Some years before" Ozeretskovskii wrote "we saw here, in Petersburg, doubled chickens, hatched from one egg, provided with all organs and united at the backs; when one of them stood on the legs, the other lied on it on the back with legs upwards in the most unnatural position. It is clear that, only the narrowness of the shell was the cause that the twin chickens, which were pressed to each other, united by the backs, similar to the united apples, which, beginning from the moment of flowering, closely adjoined each other".


1. N. Ozeretskovskii. De ovo perforata. There also, 12.1801, pp. 364-368.


Both mentioned articles of Ozeretskovskii witness that the Russian academician considered the deformity a result of the caused changes of normal development by external influences, i.e. explained their appearance epigenetically (134).

(39) K. Fr. Kielmeyer (1765-1844), is a famous German naturalist, held in great respect by the contemporaries although he hardly left published works. The most famous work of Kielmeyer was his speech "About the relation of organic powers between each other" (1793) . Kielmeyer particularly suggested the idea of powers, inherent within the living beings — irritability, sensibility and reproduction. Their combination in the form of ascending and descending rows corresponds to the stage of the individual development as well as the stage of development of all the organic world, in which, according to Kielmeyer, "the plan of nature" is found. By comparison of the stages of development with these rows, in which adult forms may exist, Kielmeyer made a conclusion, stated by him in a conversation with Gete (1797), that the higher organisms pass many stages in the process of embryonic development, in which they become lower. Shelling attached a very great importance to the ideas of Kielmeyer considering, that they usher in a new epoch in science. In fact, he found later on that before Kielmeyer a similar idea was stated by I. G. Gerder (17441863), who was a publicist, poet and philosopher. The works of Gerder, in particular, had a relation to the question discussed — "The ideas of the philosophy of the humanity history", were highly evaluated by Gete. They were popular in Russia as well and had a known effect on Karamzin, Shevryev and Maksimovich (143) .

(40) Prince A. P. Baryatinskii a personality of the Southern Secret Society and active propagandist became, towards the end of 1825, the chief of Kilchinskaya board (in Tul'chin the staff of the 2nd army was present, in which Pestel and some other members of Southern Secret Society served) (148) .


(41) Efrem Osipovich Mukhin (1766-1850) studied at the Kharkovsky College and Elisavetgradskaya Hospital School, and then at Moscow University. He was a junior assistant at the department of pathology and therapy at Moscow Medical School, and later on a professor at the university. He wrote a great number of manual books and special works (162) .

(42) Vilglin Michailovich Rikhter (1767-1822) was born in Moscow, finished Moscow University and published a doctor's dissertation in Erlangen. From 1790 he was a professor of obstetrics at Moscow University. Rikhter is the author of project "Practical obstetrical institute" and many manual books (162) .

(43) Ivan Fedorovich Wenssowitsch (1769-1811) studied at the Kharkovsky college and Moscow University secondary school, from where he joined the university, where he successively joined the faculties of philosophy, law and medicine. He finished the university and became a candidate of medicine. In 1803, he defended a doctor's dissertation in Moscow. From 1805 he was a professor of anatomy, physiology and forensic medicine. He published "journal medico-physical He died from tuberculosis at the age of 42 (163) .

(44) Yakov Kuzmich Kaidanov (1779-1885), from Kiev Ecclesiastical Academy, studied in the Petersburg medicosurgical school. After finishing there, he was sent to Vienna to study veterinary medicine. After his return back, he became a junior assistant at the Medico-surgical Academy. From 1809, he became a


   Historic Russian Embryology 1955: 1. Beginning of Embryological Investigations Lomonosov's Epoch | 2. Preformation or New Formation? | 3. Kaspar Friedrich Wolff - Theory of Epigenesis | 4. Wolff: "Theory Of Generation" | 5. Wolff: "Formation of the Intestine" | 6. Wolff's Teratological Works | 7. Wolff: "On the Special Essential Tower" | 8. Ideology of Wolff | Chapter 9. Theory of Epigenesis End of 18th Century | 10. Embryology in the Struggle of Russian Empirical Science Against Naturphilosophie | 11. Louis Tredern - Forgotten Embryologist Beginning of 19th Century | 12. Embryonic Membranes of Mammals - Ludwig Heinrich Bojanus | 13. Embryonic Layers - Kh. I. Pander | 14. Karl Maksimovich Baer | 15. Baer's - De Ovi Mammalium Et Hominis Genesi | 16. Baer's Ober Entw I Cklungsgesch I Chte Der Thiere | 17. Baer Part 1 - Chicken Development | 18. Baer Part 2 - History of Chicken Development | 19. Baer Vol 2 | 20. Third Part of the Bird Egg and Embryo Development | 21. Third Part - Development of Reptiles, Mammals, and Animals Deprived of Amnion and Yolk Sac | 22. Fourth Part - Development of Man | 23. Baer's Teratological Works and Embryological Reports in Petersburg | Chapter 24. Baer's Theoretical Views | 25. Invertebrate Embryology - A. Grube, A. D. Nordmann, N. A. Warnek, and A. Krohn

Cite this page: Hill, M.A. (2024, March 29) Embryology Book - History of embryology in Russia 1750 - 1850. Retrieved from https://embryology.med.unsw.edu.au/embryology/index.php/Book_-_History_of_embryology_in_Russia_1750_-_1850

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