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ISSN 1062-3604, Russian Journal of Developmental Biology, 2008, Vol. 38, No. 5, pp. 307–315. © Pleiades Publishing, Inc., 2008.
Original Russian Text © L.V. Belousov, 2008, published in Ontogenez, 2008, Vol. 38, No. 5, pp. 379–389.
307
More than half a century since Alexander Gavrilov-
ich Gurwitsch (1874–1954) died is a great period in the
history of Russian and international biology. Consider-
ing the rapid development of current science and its
focusing on recent findings while typically forgetting
the past, it is amazing that the personality of Alexander
Gurwitsch, probably one of the most heretical scientists
of the first half of the 20th century, is not forgotten but
rather attracts increasing interest. Some of his works
were reissued or issued for the first time 40 years after
they were written (Gurwitsch, 1991); three Interna-
tional Gurwitsch Conferences (1994, 1999, and 2004)
were held; and his representation in Internet is unusu-
ally wide for a scientist passed long ago. Recalling and
citing Gurwitsch has become a good style in Russian
publications. Would this absolutely not conceited sci-
entist be pleased with it? Let us try to impartially ana-
lyze the main in the scientific heritage of Alexander
Gurwitsch in the context of current problems of biology
(primarily, developmental biology). We will focus only
on the details of his biography directly related to typical
characters of his scientific approaches (a detailed biog-
raphy of Alexander Gurwitsch can be found in
Belousov et al., 1970).
From the beginning of his scientific carrier, Alex-
ander Gurwitsch, who graduated in the late 19th cen-
tury from the Medical Faculty of University of Munich
after specializing in histology and embryology in the
laboratory of Karl Wilhelm von Kupffer, was led by the
motivation quite unusual for biological and the more so
medical scientists. On the one hand, it was related to the
artistic gifts of the scientists (prior to becoming a med-
ical student, he tried to enroll in the Munich Academy
of Fine Arts) and his high aesthetic standards. The
beauty of mitotic figures and embryonic shapes deter-
mined his interest to cell division and embryology. It
was later reflected in the magnificent Atlas of Embryol-
ogy including 55 original color tables; it was translated
to German and Spanish and still remains a desk book
for a century (Gurwitsch, 1909)! The first versions of
the morphogenetic field theory were inspired in part by
ancient Russian churches in Rostov-Yaroslavski. Gur-
witsch’s artistic eye noticed that the outlines of the cen-
tral onion cupola in “good and truly ancient” churches
mentally extended down precisely enclose the outlines
of the side cupolas (Fig. 1a). The same pattern can be
found in the paper describing the first version of the mor-
phogenetic field theory (Gurwitsch, 1992) (Fig. 1b). The
similarity between the standards of ancient Russian
architecture (based on the concept of sobornost' (colle-
giality)) and the theory of morphogenetic field can be
not superficial (see Bishof, 2007).
On the other hand, the scientific conversations with
his relative and friend Leonid Isaakovich Mandelsh-
tam, who later became a famous physicist, had a pro-
found impact on young Gurwitsch. In particular,
Leonid Mandelshtam set forth the Einstein’s special
relativity immediately after it was published (likewise,
Moscow physicists recalled how Gurwitsch helped
Mandelshtam to tranquilize before an important exam-
ination in physics by medical means). Be that as it may,
the idea of an invariant law running through all subse-
HISTORY OF SCIENCE
“Our Standpoint Different from Common…”
(Scientific Heritage of Alexander Gurwitsch)
L. V. Belousov
Moscow State University, Moscow, 119992 Russia
e-mail: morphogenesis@yandex.ru
Received October 25, 2007; in final form, December 3, 2007
Abstract
—The work of prominent Russian biologist Alexander Gavrilovich Gurwitsch (1874–1954) on the
theory of organism development are reviewed. Alexander Gurwitsch introduced the concept of embryonic
(morphogenetic, biological, and cellular) field and proposed several revisions of it from 1912 to 1944. Although
neither of them can be considered as a final theory of development, his the persistent search for the invariant
law that allows the shape (spatial structure) to be proposed for each next developmental stage from the previous
shape is of imperishable methodological interest. Alexander Gurwitsch anticipated many ideas of the future the-
ory of self-organization. His theoretical constructions are explicit and experiment-oriented but absolutely not
esoteric. They represent a highly important and original contribution to theoretical biology and are an essential
step to further development of the ontogenetic theory.
DOI:
10.1134/S1062360408050081
Key words
: A. G. Gurwitsch, embryonic (morphogenetic) field, positional information, self-organization.
308
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quent “field” constructions of Alexander Gurwitsch
came from this source.
The time when Gurwitsch started his scientific
activity (the very beginning of the 20th century) was the
golden age of biology. Not long before, the psycholog-
ical mechanisms of fertilization have been described
and the relationship between the chromosomes and
heredity has been established. The Mendel’s Laws were
rediscovered and immediately became internationally
renowned (and Gurwitsch was the first to introduce
them to the Russian community). However, he was
most attracted by the mechanics of development—a
new trend in embryology successfully competing with
the classical descriptive approach. The founders of this
trend, Wilhelm Roux and Hans Driesch, were in the
prime of their creative lives, and Alexander Gurwitsch
soon gained their friendship and respect. “An original
thinker that sees many things differently” wrote Roux
about Gurwitsch, and later invited him to head a depart-
ment at the University of Berlin; while Driesch recog-
nized him as “a profound and ingenious researcher”
(Driesch, 1921, p. 153).
The first Gurwitsch’s study in experimental embry-
ology (which was also the first study in the field con-
ducted in Russia, in St. Petersburg), “On the Phenom-
ena of Regulation in the Protoplasm” (Gurwitsch,
1908), was widely recognized. It demonstrated high
regenerative capacity of microscopic structure in the
egg cytoplasm after centrifugation. In this work, Gur-
witsch first got in touch with the structures that go
beyond optical microscopy. Here starts the long way for
30 years to the understanding of “nonequilibrium
molecular constellations” (see below).
However, Alexander Gurwitsch left experimental
embryology immediately after this successful debut,
which perplexed well-disposed Wilhelm Roux, “Dear
colleague, why have you stopped experimenting?” The
Gurwitsch’s response was deeply reasoned. In essence,
experimenting alone cannot give rise to a theory of
development, and he considered such theory as an
invariant physical law covering maximally long devel-
opmental period. He posed a daring problem to grope
for such law, and addressed himself to the quantitative
(statistical) analysis of cell divisions and movements
during various morphogenetic processes. The range of
studied subjects was very wide: onion roots, sea urchin
eggs, and squaloid embryos (two latter subjects were
transported to St. Petersburg with express trains from
the Russian Mediterranean Biological Station at Ville-
franche-sur-mer). In all cases, Gurwitsch was inter-
ested in the relationship between elementary processes
(division, movement) at the level of individual cells and
the whole shape. Soon he concluded that these relation-
ships are far from being unambiguous. For instance,
ideally symmetrical shape of the whole results from
asymmetrical distribution of simultaneous cell divi-
sions. Thus, the whole allows wide degrees of freedom
for the elements while mildly controlling their activity.
These pioneering studies (Gurwitsch, 1910) throw a
bridge (100-year-long!) to current studies on “deter-
ministic chaos.”
At the same time, Alexander Gurwitsch attends to
the proportion between the factors of heredity and
development. As mentioned above, he was the first to
introduce the Mendel’s Laws to the Russian commu-
nity. Their mathematical expression initially impressed
him greatly. However, soon he was disappointed since
the “Mendel’s factors” (later called genes) applied only
to definitive characters; moreover, these characters
were considered mosaically and independently. Alex-
ander Gurwitsch was concerned only in the hereditary
factors that provide for forward development as a
whole. This unusual understanding is introduced in his
two conceptual publications: “Die Vererbung als Ver-
wirklichungsvorgang” (Gurwitsch, 1912) and “Der Ver-
erbungsmechanismus der Form” (Gurwitsch, 1914).
a b
Fig. 1.
Artistic prototype of the embryonic field theory. Like the outlines of the cupolas in Rostov Kremlin fit in the same envelope
(
a
), the inflorescence outlines reach the parabolic envelope during growth (
b
).
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“OUR STANDPOINT DIFFERENT FROM COMMON…” 309
These publications never mentioned genes. At the same
time, the notion of force field (kraftfeld) was first intro-
duced to biology in 1912 (Gurwitsch, 1912, p. 484).
1
The second of these papers describes a remarkable
finding: the cell axes are at some angles rather than per-
pendicular to the layer plane in the neuroepithelia in
developing telencephalon (and in the epithelium of
olfactory placodes). Moreover, if the extensions of the
cell axes are connected with a line perpendicular to all
of them, the resulting outline corresponds to the next
not yet reached developmental stage (Fig. 2a). Gur-
witsch designated such arrangement of cell axes “prog-
nostic” (of the next stage).
2
In addition, he was the first
to clearly distinguish passive and active cell move-
ments and proposed the criteria to distinguish them.
Thus, Gurwitsch laid a corner stone to morphomechan-
ics, although he considered himself a vitalist after Hans
Driesch. Now we know (see Cherdantsev, 2003) that
the prognostic orientation of the cell axes is universal
for all epithelial morphogeneses without exception (the
current (morphomechanical) interpretation of this phe-
nomenon (Fig. 2b) is discussed below). For Gurwitsch,
this indicated that the space surrounding the cell layer
contained a “force surface” that rotated the cell axes
and colocalized with the outline of the subsequent
developmental stage (at a definite not very short time
distance from this stage). Gurwitsch named this factor
a “dynamically preformed morph.” Several years later,
the notion of dynamically preformed morph was
extended to the development of fungi and Compositae
1
In this publication, the notion of force field was mentioned in
passing, and it was missing from the subsequent publication
(Gurwitsch, 1914) (it was replaced with “dynamically preformed
morph”). Only in 1922, “embryonic field” was included in the
paper title (Gurwitsch, 1922). This gave grounds to question his
priority in introducing the field notion to biology (see commen-
tary to Beloussov, 1997 by John Opitz and Scott Gilbert). At the
same time, it is clear in the context of the papers published in
1912 and 1914 that Gurwitsch has elaborated the field notion
then.
2
Actually, Gurwitsch wrote about the axes of cell nuclei rather
than of cells. Here, the histological dyes that stained the nuclei
but not cell walls played a bad trick with him, and he erroneously
concluded that the cell nuclei rather than the whole cells are
reoriented. This could be crucial for the later views of Alexander
Gurwitsch on the nuclei as a field source.
plants in the first publication with the words “embry-
onic field” in the title (Gurwitsch, 1922). Later, the
development of the embryonic field theory decelerated
again, this time due to intense experimental studies of
ultralow mitogenic radiation, which he discovered in
the same 1922. The next step was made by Gurwitsch’s
disciple, Andrei Anikin, who proposed the invariant law
of nucleus deformation in chondrocytes during the
development of finger bones in newt (Anikin, 1929).
Most importantly, this work replaced the concept of
dynamically preformed morph with a more physicalis-
tic principle of field action from a point source.
Strangely enough, the tragic events of the Second
World War gave an impetus to the development of the
field theory. Having no opportunity for experimental
work during the siege of Leningrad in October–Decem-
ber 1941 under artillery fire by monstrous descendants
of his German teachers and colleagues, Gurwitsch rap-
idly outlines a new version of his theory not being sure
that anyone will know about it. In excitement we now
turn the pages of the hastily self-bound notebook (Fig.
3). The record of November 5, after an intense night air
bombardment when several firebombs were dropped in
the area of the All-Union Institute of Experimental
Medicine, where Gurwitsch lived and worked, says
nothing about it. It contains a detailed analysis of the
rotation of mitotic spindles during early cleavage in
ascarid—the scientist tries to elaborate algorithms of
the field from these processes. However, the theoretic
outlines on November 13 are interrupted with a short
entry “Situation deteriorating; progressing malnutri-
tion,” and then “Returning to equilibrium structures…”
Luckily, the scientist was saved, and two years later
he “not without hesitation” publishes a new version of
his theory (Gurwitsch, 1944). Let us try to understand
his main intention using this theory and what distin-
guishes it from other concepts aimed to solve similar
problems.
As Gurwitsch repeatedly mentioned, the starting
point of all his field concepts was the famous Driesch’s
Law: The prospective fate of an embryonic part is a
function of its position within a whole. However, what
was the final point in experimental work for Driesch
(his further interests were purely philosophic), was
only the starting point for Gurwitsch. He realized that a
a b
Fig. 2.
Comparison of the design of dynamically preformed morph (
a
; Gurwitsch, 1914) with the experiment on neuroepithelium
separation in brown frog embryo at the stage of early neurula (
b
);
)
, separation direction.
310
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general theory of embryonic development can be built
on the basis of this law. However, this required that the
meaning of “fate” should be strictly defined. What is
the effect of the “factor of the whole” on the parts? Can
the Driesch’s Law be revised to become applicable not
only to isolated experimental conditions but also to
common events during normal development? In order
to find the most specific and empirically testable
answers, Alexander Gurwitsch first proposed to con-
sider the fate of a particular part as a set of vectors of its
morphogenetic movements. These should be the target
for the effect of the whole. The latter interpretation var-
ied most in Gurwitsch’s concepts. Initially, the whole
was understood as a dynamically preformed morph
suspended in space or, in the Anikin’s work, as a point
source unrelated to material particles; while in the the-
ory of 1944, it arose from the vector fields of individual
embryonic cells, and cell nuclei (according to Gur-
witsch, chromatin molecules during their synthesis)
were the field sources. Such splitting of the whole into
a set of individual sources disappointed not numerous
advocates of his previous concepts, who claimed that
Gurwitsch has abandoned the idea of wholeness. Gur-
witsch completely disagreed (Gurwitsch, 1947) and, in
our view, he was absolutely right. First, the properties
of the whole were retained in the new theory, since the
geometry
of an embryo or rudiment played the key role.
Second, the whole no more emerged from nowhere and
was introduced ad hoc for an arbitrary period of space
and time, as in the concept of dynamically preformed
morph. Conversely, a concerted evolution of the whole
and parts appeared: the changes in cell positions
induced by the field of the whole have a reciprocal
effect on the whole and vice versa—the embryonic tis-
sue started self-coordinated movement underlain by
feedback relationships. On these grounds, the theory of
Fig. 3.
Notes of Alexander Gurwitsch in November 1941. Upper part, a diary record: “Interruption until November 11—The chapter
with systemic presentation of the field was written during this period—then a full interruption: departure of children November 5
−
7.
Situation deteriorating; progressing malnutrition.” Lower part, three fragments of the above-mentioned chapter. Note that it was
written in German—despite the deteriorating situation, it was easier for Gurwitsch to write scientific texts in this language.
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“OUR STANDPOINT DIFFERENT FROM COMMON…” 311
1944 demonstrated that, basically,
forms of subsequent
developmental stages can be deduced from the previous
forms, i.e., the causal chains of morphogenesis can be
closed at the macromorphological level to a good
approximation
without repeated addressing to the pro-
cesses at different levels (e.g., molecular). On the one
hand, this gave an impetus to the ideas of pre-Darwin
rational (idealistic) morphology about immanent “gen-
erative rules of morphogenesis” that have to be found
(see Webster and Goodwin, 1982). On the other hand,
the theory of 1944 anticipated a number of fundamental
principles of the later theory of self-organization. In
modern terms, the main idea of the theory of 1944 is an
attempt of a closed description of morphogenesis based
on the self-coordinated field.
In his attempts to close the interpretation/descrip-
tion of development at the macromorphological level,
Gurwitsch never ignored the role of molecular (submi-
croscopic) processes; conversely, they increasingly
attracted his special attention in the late period of his
creative activity. At the same time, he stated that “the
utilization pattern of submicroscopic structures… can-
not be deduced from their own parameters” (Gur-
witsch, 1991, p. 105).
It is apt here to mention one more principal state-
ment, the first words of which were included in the title,
“our standpoint, different from a common one, is that a
really adequate analysis of any given developmental
stage should conclude its transition to the next stage”
(Gurwitsch, 1991, p. 124). Why he considered this
standpoint uncommon? Didn’t mechanics of develop-
ment declare finding the grounds of an embryo at a par-
ticular developmental stage for the transition to the next
stage as one of major objectives? Are not, e.g., the
inductive properties of chordomesoderm discovered by
Hans Spemann the grounds for inevitable transition
from gastrula to neurula? Here lies the principal dis-
tinction between the ideology of developmental
mechanics, popular in scientific community, and the
ideology of Gurwitsch.
The ideology of developmental mechanics and
nearly all current biology is based on the notion of fac-
tors
specific
for each individual developmental step and
process. In this case, the research methodology consists
in studying these factors separately and by dissecting
the studied subject, and the ultimate research goal is a
maximally complete list of such factors and their
effects. The problem of a law common for this list is not
even raised. This approach is unacceptable for Gur-
witsch: “a problem is not an individual phenomenon
but rather their long continuous sequence or chain. Let
us assume for a moment that individual mechanisms
can be proposed for each of conventional elements of
this sequence A, B, and C; e.g., process A is reduced to
swelling, process B is reduced to chemical reaction,
etc. However, the interest to these mechanisms recedes
into the background of the problem why this regular
sequence of, apparently, quite unlike processes exists”
(Gurwitsch, 1944, p. 8).
Gurwitsch considered the reduction of such a
sequence to a list of purely specific factors equal to the
denial of scientific explanation. In addition to his own
arguments appealing to “the nature of our sense” (Gur-
witsch, 1944, p. 8), let us present purely biological
arguments partially known in his time but substantially
amplified later.
First, the analytical (separating) search for new spe-
cific factors leads to the dead-end of preformism: a par-
ticular factor should be controlled by another also spe-
cific (at least in terms of its spatiotemporal localization)
factor and so on indefinitely. This is called “irreducible
complexity” in modern science (see Behe, 1996). At the
same time, we consider the establishing conclusion of
current molecular and cell biology about
the absence of
an ambiguous relationships between molecular factors
(inducing substances, genes, or signaling cascades)
of
particular developmental processes and the proper
processes considered at the cellular or supracellular
levels.
Manuals of cell molecular biology and develop-
mental biology are crowded with examples of this kind.
Stated differently, even exhaustive data on the molecu-
lar processes in a given embryonic rudiment do not
allow us to predict even the next developmental stage;
moreover, we cannot identify the rudiment (cf. Gur-
witsch’s “Our hope that the elucidation of submicro-
scopic structures in the egg cell is the way to understand
(predict) at least the next in time vital form is not real-
izing;” Gurwitsch, 1991, p. 105). The great achieve-
ments in molecular and cell biology in the recent
decades have not brought us closer to the answer to a
naive question: why form A is replaced with form B and
so forth during body development? This question is
ignored and as though does not exist. Instead, current
molecular biology is (reasonably) proud of identifying
many
universal
signaling cascades, gene circuits, etc.
that were largely conserved over huge evolutionary dis-
tances. This clearly outstanding achievement is useless
to solve the main ontogenetic problem: why stage
(form) A transmits to stage (form) B. Universal molec-
ular mechanisms are
tools
of morphogenesis (Driesch
called them
mittel
) that are required for particular pro-
cesses; however, they cannot determine what and when
they will effectuate. The Gurwitsch’s theory of 1944 is
the only constructive attempt to solve this naive ontoge-
netic problem in modern science. Abandoning this
attempt would mean that ontogeny has no intrinsic
dynamics and corresponds to execution of a set of
“instructions,” the sequence of which is self-given and
defies explanation. Is biology doomed to uncondition-
ally accept this “instructivistic” viewpoint or the gen-
eral laws of development still exist? For Gurwitsch,
accepting the first viewpoint would mean that “a given
sequence of events is scientifically inaccessible” (Gur-
witsch, 1944, p. 9). The only scientific solution of this
problem for him was “the resolution (of a given cycle
of events, L.B.) into a definite not very high number of
312
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stages, each of which could be presented as a monotone
function of specific conditions prescribed in the initial
conditions with a single variable in the form of time,
travel, etc.” (Gurwitsch, 1944, p. 9).
Unfortunately, the Gurwitsch’s attempt to propose
the general developmental laws of this kind was not
entirely successful. The theory of 1944 had one more
advantage that became fatal for it—it is so specific that
can be easily falsified (which hardly applies to most
other biological theories). The point is that Gurwitsch
could not avoid a universal algorithm of field source
effect on cells. Based on the data available to him
(largely borrowed from the publication of Anikin), he
proposed that this algorithm consists in cell repulsion
from the field sources inversely with the square of the
distance. At first, this assumption was confirmed for
some rudiment types. I still remember my astonishment
40 years ago, when I applied not complicated repulsion
constructions of Gurwitsch to the rudiments in
hydroids (he knew nothing about them) to yield an out-
line of the next developmental stage (Belousov, 1968a,
1968b)! Several years of work were required to demon-
strate that the observed sequence of forms does not
require the field action through the cell-free space. It is
entirely derived from strictly recorded periodic pulses
of internal pressure in the cell layer, which have
osmotic nature (Beloussov et al., 1989; Beloussov and
Grabovsky, 2003). In addition, the main morphogenetic
movements in vertebrate embryos, radial and conver-
gent cell intercalation, directly contradict the repulsion
hypothesis, since cells conversely approach each other.
However, before returning a verdict to the theory of
1944, let us compare it to the concept relatively popular
nowadays, which also used the Driesch’s Law as the
starting point—the positional information concept
(Wolpert, 1969, 1996). The positional information
relies on the notion that the differentiation fate of an
embryonic cell depends on its position in a concentra-
tion gradient of certain compound, a morphogen. The
gradient lies between preset “special points” commonly
called sources and stocks. Each individual cell is
thought to read its position in a gradient and interpret it,
i.e., to choose its differentiation fate according to its
specific genome.
Indeed, there is a relationship between the direction
of cell differentiation and the concentration of certain
factors binding protein receptors of the cell membrane,
cytoplasm, or nucleus (inducers, retinoic acid, etc.) (see
Lander, 2007). However, even superficial analysis of
real cell responses to concentration gradients points to
nonlinear relationships typical of the whole system.
This primarily applies to the pronounced threshold dis-
tribution pattern of various cell types along the initially
monotone gradient. According to the theory of self-
organization, the role of positional factors in such (e.g.,
metameric) systems is reduced to establishing marginal
conditions, while the parameters specific for the system
as a whole become crucial for the “layout.” The essen-
tially mosaic concept of positional information does
not assume them. However, it deserves the most serious
criticism in general terms: what it explains in develop-
mental processes? The problem of deducing the result
of development from positional information is just not
raised; instead, one is referred to enigmatic interpreta-
tion of positional information by the genome. This con-
cept bears no relation to morphogenesis per se—it
explains neither the formation nor possible effect of the
forms on cell differentiation assuming very simple con-
cept of development as a direct transformation of a
chemical gradient into a differentiation layout. More-
over, one can easily demonstrate that considering posi-
tional information as a set of gradients directly conflicts
the embryonic regulation processes, which were the
target for the Driesch’s Law. Indeed, any preset gradi-
ents linked to specific sites in the embryo can occupy
various positions geometrically nonhomologous to the
normal ones after the removal, addition, or mixing the
embryonic material. As a result, the spatial distribution
of the positional information and, hence, the embryonic
structure will be inevitably disturbed (discussed in
detail in Belousov, 1987; Beloussov et al., 1997;
Beloussov, 1998). The only way to maintain the invari-
ant structure of the whole after experimental actions
mentioned above is to reject special reference points for
positional information and consider that each part
determines its fate from the position relative to the sets
of all other furthermore
equivalent
parts, the mutual
position of which plays the key role. This returns us to
the theory of Gurwitsch.
In our opinion, the above considerations suffice to
state that the concept of positional information cannot
pretend to be even an outline of the developmental the-
ory. As a matter of fact, it does not pretend. Many pub-
lications use it as a noncommittal excuse to obviate
inconvenient problems. In the intrinsic structure, the
Gurwitsch’s theory of 1944 is much more consistent
and specific. However, it also directly conflicts some
firmly established facts as shown above. Can these con-
flicts be resolved while keeping the ideology of the
Gurwitsch’s theory—specific description of the inter-
action of whole and parts and the “form from form”
thesis?
Our research group at the Department of Embryol-
ogy at Moscow State University pursued this line from
1970s and has reached the following conclusions. At
least morphogenesis per se, i.e., a set of morphogenetic
cell movements (and possibly major types of cell differ-
entiation, since they are related to morphogenesis), can
be presented as a sequence of chains that can be
deduced from each other if considered as
active mech-
anochemical responses to previous mechanical strain
in embryonic tissues. An algorithm of these responses
was proposed—the hypothesis of “hyperrestoration of
mechanical strains” (Belousov and Mittental’, 1992;
Beloussov et al., 1994, 2006; Beloussov, 1998). Thus,
the “form from form” principle is replaced with the
“form from mechanical strain field” principle. In terms
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“OUR STANDPOINT DIFFERENT FROM COMMON…” 313
of the succinct statement of D’Arcy Thompson “the
form of an object is a diagram of forces” (D’Arcy
Thompson, 2000), this replacement seems admissible:
there are definite and clearly formulable relationships
between the primordial form and the field of its
mechanical strains, although these notions are clearly
not identical (the same form can be strained or relaxed).
At the same time, the positional information notion
becomes not self-contained, since not the geographical
coordinates but rather the morphomechanical history
(memory) over a short but definite previous period of
development is important for the fate of an embryonic
element. Stated differently, not the position alone but a
combination of the position and the nearest morphome-
chanical history determine the fate for a given embry-
onic part.
The morphomechanical grounds make it possible to
reconstruct (model) quite long morphogenetic periods.
In this case, its universal archetypes are outlined, on the
one hand, and the species diversity reproduction
becomes possible, on the other hand (Beloussov et al.,
2006). The latter is mediated by changing variables,
while the universal algorithm remains invariant. The
variables can be linked to both genetic and epigenetic
events.
Within the frames of the morphogenetic approach,
some old problems of the field theory find unexpectedly
simple solutions. For instance, the dynamically pre-
formed morph exactly coincides with the surface, to
which momentarily (within several seconds) the ini-
tially strained epithelial layer surface is relaxed after
the mechanical strain is relieved (Fig. 2b).
Taking into account all above considerations, we
can regard the morphogenetic approach as a natural
development of the Gurwitsch’s theory of morphoge-
netic fields.
In conclusion, let us discuss Gurwitsch’s concepts
concerning the processes at the molecular level. As
mentioned above, Gurwitsch was the first to notice the
morphogenetic importance of processes that go beyond
optical microscopy in his work on egg cell centrifuga-
tion. His interest to molecular processes was amplified
during his intense study of ultralow radiation. He par-
ticularly emphasized “degradational radiation,” short-
term explosions of photon emission after reversible
damage (temperature, mechanical, etc.) (Gurwitsch and
Gurwitsch, 1945). According to Gurwitsch, these
explosions indicated the presence of nonequilibrium
(requiring continuous energy influx) molecular struc-
tures integrating energy potentials in the range from the
basal metabolic level (around 0.5 eV) to 5 eV (the
energy of UV photons). Gurwitsch called these struc-
tures nonequilibrium molecular constellations. Note
that this notion completely corresponded to the notion
of dissipative structures that appeared in the theory of
self-organization several decades later. How can non-
equilibrium molecular constellations emerge and self-
maintain? Gurwitsch believed that this is mediated by
to a certain extent
vectorized
(i.e. different from cha-
otic/diffusion) movement of small molecules and
deformation of large molecules in the cytoplasm. He
attributed the vectorization of molecular movements to
the field factor. According to his statement, the mole-
cules with internal energy are the field targets, the func-
tion of which is to transform a part of this energy into
directional kinetic energy (oriented along the field vec-
tors) (Gurwitsch, 1944, 1991).
While considering these central statements of the
Gurwitsch’s theory, one should bear in mind that they
were proposed in 1930s–1940s, when almost nothing
was known about cytoplasmic structures and the more
so about the movement of cytoplasm components.
Nowadays, the interest of Gurwitsch to vector move-
ments in the cytoplasm looks prophetic. More strict
parallels can be found, e.g., the field function as a trans-
former of nondirectional energy into directional energy
exactly corresponds to the current views on growing
microtubules as Brownian ratchets (see Peskin et al.,
1993). However, very diverse movements of subcellu-
lar particles and molecules (in different directions) can
hardly be related to some common field, particularly,
originating from cell nuclei, in the light of the currently
available data. It is common knowledge that vector
movements are possible in nucleus-free cell fragments
and that they are due to the polymerization and depoly-
merization of cytoskeletal structures and activity of
special molecular motors. In this case, there is no way
to avoid the specificity and to reduce all intracellular
movements to the effect of a global factor.
As mentioned above, the notion of nonequilibrium
molecular constellations introduced by Gurwitsch is a
precise anticipation of the notion of dissipative struc-
tures central in the theory of self-organization. Never-
theless, the causal chains in the Gurwitsch’s field the-
ory and in the theory of self-organization are contrary.
According to Gurwitsch, the maintenance of nonequi-
librium molecular order
requires
a field that is
external
relative to it. Conversely, the molecular nonequilibrium
creates
such self-coordinated field according to the
self-organization theory. Elementary examples include
Benard structures, Beloussov-Zhabotinsky structures,
etc. We do not mean that Gurwitsch was wrong and the
theory of self-organization was right concerning the
self-organization, but rather wish to draw attention to
two contrary theoretical approaches based on the same
experimental data.
At the same time, one cannot but admit that the Gur-
witsch’s creative signature, the urge to minimize spe-
cific components of biological processes and to boot in
the foreground the global invariant factors, proved
more efficient for morphogenetic processes at the
whole embryo level than for intracellular processes. In
the latter case, the molecular specificity and short-range
molecular interaction clearly play a very significant
role. Nonetheless, this does not mean that we are
doomed to purely additive description of cell activity
314
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Vol. 38
No. 5
2008
BELOUSOV
and that problems such as “cell as a functioning whole”
or “genome as a whole” (giant body, see Ingber, 2005)
are senseless. The same problem exists here as in the
case of morphogenesis: are the sequences of molecular
intracellular processes closed at the level of specific
intermolecular interactions or they are influenced by
less specific processes at higher organization levels (in
this case, represented by the cell shape, architecture of
its contacts and of course the fields of mechanical
strain)? Some recent data (see Huang and Ingber, 2000;
Ingber, 2005) clearly support the latter possibility.
Moreover, shocking statements concerning “molecular
vitalism” can be found (Kirschner et al., 2000). Any-
way, application of the field principle to intramolecular
processes requires considerable work and fine
approaches.
In summary of this rather cursory review of Alex-
ander Gurwitsch’s theoretical concepts, it should be
stressed that they were always specific and testable and
targeted to direct application in empirical studies. Nat-
urally, esoteric deviations were absolutely unaccept-
able for Gurwitsch. In this context, it seems highly
unlikely that he would be glad with the image of vague
and almost mystical thinker sometimes arrogated to
him nowadays. He would strongly object the equation
of field factors and ultralow radiations, which is not
uncommon. In our view, the methodology is the stron-
gest point in the Gurwitsch’s concepts, i.e., the manner
to define problems and tochoose approaches to them
rather than the results. In this context, the knowledge of
scientific heritage of Alexander Gurwitsch, that has
deep historical roots and is turned to future, is not only
beneficial but also necessary for anyone who wants to
see biology a fundamental science.
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