University of Virginia Library

RECAPITULATION

The theory of recapitulation, designated the Bio-
genetic Law by Ernst Heinrich Haeckel in 1872
(1872a), stated in brief that ontogeny, the development
of the individual, recapitulates phylogeny, the evolu-
tionary history of the stock to which the individual
belongs. The fact that it is still appropriate to discuss
it in the 1970's is a tribute to the viability of an idea
that was only tenuously supported by scientific evi-
dence when put to the test of validity.

The concept attained its greatest vigor after the
publication of Darwin's Origin of Species (1859). When
at the end of the nineteenth century a new science,
experimental embryology, demonstrated that particu-
lar developmental processes might be analyzed in
terms of more proximate events, embryological interest
in the concept waned. When in the twentieth century
the science of genetics developed and coalesced with
the study of evolution, investigators in these fields also
lost interest in recapitulation, yet as late as the 1950's
a prominent embryologist stated categorically in a
textbook of embryology: “Today there remains no
reasonable doubt about the fundamental fact that
developing vertebrates pass through series of stages
which in general recapitulate the evolutionary
progression” (Witschi, 1956). He added some qualifi-
cations, and such a statement is rare in middle and
late twentieth-century treatises, but the dogma lingers.

Before Darwin crystallized the idea that all orga-
nisms are related genealogically, the concept that or-
ganisms could be ranked from low to high on a Scale
of Beings imbued the thought of philosophers and
scientists. When this theory carried authority, similari-
ties between embryos of one form and adults of another
were described in generalizations that lacked evolu-
tionary connotations. Some historians trace such con-
cepts back to Aristotle, who believed that man during
his development had first the soul of a plant, then that
of an animal, before finally acquiring his rational soul.
This was a metaphysical concept rather than a biologi-
cal one, as perhaps also was William Harvey's expres-
sion of a parallel between embryos and animals
ascending the Scale of Beings. “Nature,” wrote Harvey,
“ever perfect and divine, doing nothing in vain, has
neither given a heart where it was not required, nor
produced one before its office became necessary; but
by the same stages in the development of every animal,
passing through the constitutions of all, as I may say
(ovum, worm, foetus) it acquires perfection in each”
(Harvey, 1628).

The concept, metaphysical or otherwise, was alive
apart from biological thought during the years when
it began to attract scientific attention. For instance,
William Blake, describing the development of Orc in
the womb of Enitharmon, put it thus (Blake, 1794):

It was during Blake's time, however, that the concept
began to acquire scientific status. Kohlbrugge (1911)
presented a list of over seventy-two writers, beginning
with Goethe and Autenrieth, who expressed ideas of
parallelism or recapitulation between 1797 and 1866,
the year that Haeckel formulated the recapitulation
theory in a few catchwords that were to become a
slogan.

Johann Friedrich Meckel was one of the most im-
portant of many who expounded laws of parallelism
during the nineteenth century. He wrote in his treatise
on comparative anatomy, when comparative anatomy
was becoming a dominant biological discipline: “An
embryo of higher animals passes through a number

of stages before it attains its complete development;
it is to be demonstrated here that these different
stages correspond to those at which lower animals
are arrested throughout their whole lives” (Meckel,
1821).

Meckel's generalization was soon to be disputed by
a master as great as himself. Within five years, Karl
Ernst von Baer wrote in an article that: “It has been
concluded by a bold generalization from a few analo-
gies, that the higher animals run in the course of their
development through the lower animal grades, and
sometimes tacitly and sometimes expressly they have
been supposed to take their way through all forms.
We hold this to be not only untrue, but also impossible”
(von Baer, 1827). Von Baer's treatise Ueber Ent-
wicklungsgeschichte der Thiere (On the Embryology
of Animals), which established comparative embryol-
ogy as a discipline that was soon to supplant compara-
tive anatomy as the most important zoological science,
devoted one of its most important sections to a refuta-
tion of the law of parallelism. In it he deduced on the
basis of his own careful observations what he called
“the law of individual development, 1. That the more
general characters of a large group of animals appear
earlier in their embryos than the more special charac-
ters.... 2. From the most general forms the less gen-
eral are developed, and so on, until finally the most
special arises.... 3. Every embryo of a given animal
form, instead of passing through the other forms, rather
becomes separated from them.... Fundamentally,
therefore, the embryo of a higher form never resembles
any other form, but only its embryo” (von Baer, 1828).
As one recent authoritative critic of recapitulation
theory has paraphrased it: “Instead... of passing
through the adults of other stages of other animals
during its ontogeny, a developing animal moves away
from them, and the ontogenetic stages do not run
parallel to the sequence of forms of the scale of beings”
(de Beer, 1958).

Ironically, the law of recapitulation is often
attributed to von Baer in spite of his vehement denial
of the validity of the law of parallelism. This may be
accounted for by the fact that Darwin, in the last four
editions of the Origin of Species quoted a statement
by von Baer to the effect that: “'Embryos of birds,
lizards, and snakes, probably also of chelonia are in
their earliest states exceeding like one another'”
(Darwin, 1861, 1872), and Darwin himself in the final
edition withdrew some earlier reservations and stated
categorically that: “Several... highly competent
judges insist that ancient animals resemble to a certain
extent the embryos of recent animals belonging to the
same classes; and that the geological succession of
extinct forms is nearly parallel with the embryological
development of existing forms. This view accords
admirably well with our theory” (Darwin, 1872).

Haeckel read the first edition of the Origin of Species
in German translation in 1860, and, carried away by
it, became Darwin's champion and popularizer in
Germany. He went far beyond Darwin, and sometimes
far beyond fact—he was accused of falsifying pictorial
illustrations to support his theories (His, 1874)—in
attempting to synthesize all that was known of nature
in a single scheme based on evolutionary theory.

In his first and possibly most important general work,
Generelle Morphologie der Organismen (1866; General
Morphology of Organisms), Haeckel began to relate
ontogeny to phylogeny (words that he invented):
“Ontogenesis or the development of the organic indi-
vidual, as the series of changes in form which every
individual passes during the whole period of its indi-
vidual existence, is immediately conditioned [bedingt]
by the phylogenesis or the development of the organic
stock (Phylon) to which it belongs.... Ontogenesis
is the short and rapid recapitulation of phylogenesis,
caused by the physiological functions of inheritance
(reproduction) and adaptation (nourishment).” Already
in this first of his generalizations Haeckel implied
through the use of the verb bedingen a causal rela-
tionship between ontogenesis and phylogenesis. When
in the late 1860's embryologists such as Wilhelm His
(whom Haeckel singled out for particular anathema)
attempted to explain the development of particular
organs in terms of such mechanical factors as the fold-
ing of embryonic layers caused by unequal growth,
Haeckel (1874) replied by insisting that: “Phylogenesis
is the mechanical cause of ontogenesis,” and would
tolerate no opposition.

Haeckel was wary of claiming absolute applicability
for the biogenetic law and often wrote of recapitula-
tion as “short,” “rapid,” or “abbreviated.” It was
known by the 1860's that various nutritive, respiratory,
and excretory organs (yolk-sac, allantois) characteristic
of embryos are lacking in all adults. Haeckel (1875)
took cognizance of such facts by classifying as
palingenetic the embryonic processes that are trans-
mitted by heredity from ancestral forms, in contrast
to coenogenetic ones which appear through adaptation
to the needs of embryonic or larval life.

He felt that in spite of these limitations, the clues
given by embryos concerning the ancestry of their line
were intelligible, and he constructed elaborate geneal-
ogies on the basis of the study of embryonic form.
When there were gaps in his postulated schemes of
evolutionary relationships, he was not above inventing
animals to fill them. Leaning heavily on the nine-
teenth-century concept of fixity of origin and differen-
tiation of the embryonic germ-layers (ectoderm,

mesoderm, endoderm), he postulated a two-layered
organism named Gastraea as the ancestor of all many-
celled animals (Haeckel, 1872b).

One of the most fruitful results of his forceful expo-
sition and vehement defense of the biogenetic law was
that others, like himself, studied embryos in order to
ascertain the ancestry of various species, and whether
or not the premisses from which these studies devel-
oped were sound, a great corpus of embryological
knowledge was established.

A number of factors contributed, during the late
nineteenth and early twentieth centuries, toward the
decline of embryologists' interest in recapitulation. As
biology in general concentrated less on evolution and
more on experimental physiology, embryologists be-
came more interested in experimental than in theoret-
ical approaches to the study of developmental me-
chanics. The comparability and fixity of the
germ-layers in different animals were challenged by
experimental evidence. As genetic data accumulated,
coenogenesis became less acceptable as an explanation
of deviations from strict recapitulation, since it implied
the inheritance of acquired characters, the Lamarckian
theory, which was unacceptable to the new genetics.
In other ways also, it became clear that coenogenesis
could not suffice to explain deviations from the rule.
In some groups of insects, species unlike in adult form
have similar larvae; in others, unlike larvae develop
into similar adults (de Beer, 1958). Embryos of one
order of molluscs, the cephalopods (squid, cuttlefish,
octopus) have a unique type of development; they pass
through no stages comparable to those of other
molluscs and no other molluscs pass through stages
comparable to those of cephalopods. In such organisms,
genetic changes have been introduced early in the life
history; Haeckel's concept called for them to occur
at the end of the embryonic period.

The occurrence of mutation in genes that act early
in development is a primary factor leading to the
divergencies of development emphasized nearly a cen-
tury and a half ago by von Baer. Since differences in
development lead to differences in adults, modification
of ontogeny produces new raw materials for natural
selection to work upon. From this point of view
(Garstang, 1922), most biologists of the late twentieth
century consider ontogeny the cause of phylogeny,
rather than the reverse, on the few occasions when
they consider such relationships at all.

Haeckel's ideas about recapitulation appealed not
only to biologists, but also to many others who became
aware of them because of the popularity of the books
he addressed to general readers. Like many aspects of
Darwinian theory, Haeckel's concept of recapitulation
had its impact on anthropology, criminology, political
theory, literature, psychology, and even history itself.
An example of its influence on cultural history is pro-
vided by an organic analogy used by John W. Draper:
“The march of individual existence shadows forth the
march of race-existence, being, indeed, its repre-
sentative on a little scale.... A national type pursues
its way physically and intellectually through changes
and developments answering to those of the individual,
and being represented by Infancy, Childhood, Youth,
Manhood, Old Age, and Death respectively.... Na-
tions must undergo obliteration as do the transitional
forms offered by the animal series. There is no more
an immortality for them than there is an immobility
for an embryo in any one of the manifold forms passed
through in its progress of development” (Draper, 1876).

Draper's thought was of considerable significance in
its influence on the development of intellectual history.
At a much more popular level, the transfer of the idea
of recapitulation into general thinking is exemplified
in its expression in a book on child care that was a
handbook in many thousands of American homes in
the mid-twentieth century. “Each child,” wrote
Benjamin Spock, “as he develops is retracing the whole
history of mankind, physically and spiritually, step by
step. A baby starts off in the womb as a single tiny
cell, just the way the first living thing appeared in the
ocean. Weeks later, as he lies in the amniotic fluid in
the womb, he has gills like a fish” (Spock, 1968).

The concept of recapitulation was especially influ-
ential in psychology, in which Haeckel himself was
particularly interested. During the early part of the
twentieth century it was incorporated into psychology
by Carl Gustav Jung, who adopted the theory in sup-
port of his own ideas about the “race unconscious”:

(Jung, 1916; trans. 1927).

Jung first developed his idea of the race or collective
unconscious through the consideration of dreams, and
confirmed it by his analogy to the development of the
body. Analogy between the development of mind and
body, on the basis of recapitulation, was made as early
as 1866 by Haeckel in his General Morphology: “The

psychic life of man obeys completely the same laws
as the psychic life of other animals.... Like all com-
plicated phenomena in higher organisms, so the mind,
as the most complicated and highest function of all
... can be understood only by comparing it with
simpler and less complete phenomena of the same sort
in lower animals and by following its gradual develop-
ment step by step. We must return here not only to
the biontic but also to phyletic development.”

Wilhelm Preyer, Professor of Physiology at Jena
during twenty-seven of Haeckel's years in the same
university, extended Haeckel's ideas to child psychol-
ogy. “The mind of the new-born child,” he wrote in
the preface to the first edition of Die Seele des Kindes
(1882; The Mind of the Child), “does not resemble a
tabula rasa upon which the senses first write their
impressions,... but the tablet is already written upon
before birth, with many illegible, even unrecognizable
and invisible marks, traces of the imprint of countless
sense impressions of long past generations.” And Preyer
went further than Haeckel to extend the concept to
more specific aspects of mentality, including memory,
as Jung also was to do later. “I call [personal memory],”
wrote Preyer, “the memory formed by means of indi-
vidual impressions, occurrences, experience, in contrast
with phyletic memory, or instinct, the memory of the
race, which results from the inheritance of the traces
of individual experiences of ancestors.”

Preyer may not have been the only writer in his
time to believe that the life of the psyche obeys the
same laws in its development as the life of the body,
according to the law of recapitulation, but his Mind
of the Child was an influential book, a pioneering
analysis of child psychology. It may well have been
read by Jung during his formative years, and Jung, of
course, like so many of his contemporaries, may also
have read books by Haeckel. Be that as it may, it was
of critical importance for the subsequent history of
embryology itself that The Mind of the Child was read
by Hans Spemann, who was to perform in the first
quarter of the twentieth century the experiments on
embryonic induction that were so fateful for the subse-
quent development of experimental embryology.
Spemann stated specifically in his autobiography that
he was first drawn into biology as a result of reading
The Mind of the Child. It is ironic that the same ideas
that so strongly influenced psychology through Jung's
acceptance of the recapitulation doctrine were also to
lead up to the experimental analysis which finally
destroyed the concept for biology proper.

BIBLIOGRAPHY

K. E. von Baer, “Beiträge zur Kenntnis der niedern
Thiere,” Nova Acta Physico-medica Academiae Caesareae
Leopoldino-Carolinae Naturae Curiosorum, XIII, Part 2
(1827), 525-762. Part was translated by T. H. Huxley in
Scientific Memoirs, ed. A. Henfrey and T. H. Huxley
(London, 1853); the quotation is from Huxley's translation,
p. 184; idem, Ueber Entwicklungsgeschichte der Tiere, 2 vols.
(Königsberg, 1828; 1837), I, 224; the quotation is from
Huxley's translation, Scientific Memoirs, op. cit., p. 214.
G. R. de Beer, Embryos and Ancestors, 3rd ed. (Oxford, 1958),
is the most recent full critique of recapitulation theory. W.
Blake, The First Book of Urizen, Etched 1794, in Poetry and
Prose of William Blake, ed. G. Keynes (London, 1927), p.
254. C. R. Darwin, Origin of Species..., 3rd ed. (London,
1861), pp. 470-71. Darwin's quotation from von Baer is from
Huxley's translation, Scientific Memoirs, op. cit., p. 210; The
Origin of Species..., 6th ed. (London, 1872). The quota-
tion from the 6th ed. is taken from the reprint (New York,
1902), II, 120. J. W. Draper, History of the Intellectual
Development of Europe, rev. ed., 2 vols. (New York, 1876),
I, 12, 14, 17-18. W. Garstang, “The Theory of Recapitula-
tion. A Critical Restatement of the Biogenetic Law,” Jour-
nal of the Linnaean Society of London, Zoology, 35 (1922),
81-101. E. H. Haeckel, “Die Gastraeatheorie, die physio-
logische Klassifikation des Tierreichs und die Homologie der
Keimblätter,” Je naische Zeitschrift für Naturwissenschaften,
8 (1874), 6; idem, “Die Gastrula und die Eifurchung der
Tiere,” Je naische Zeitschrift für Naturwissenschaften, 9
(1875), 402-19; idem, Generelle Morphologie der Orga-
nismen, 2 vols. (Berlin, 1866), II, 300, 344; Die Kalk-
schwämme, Biologie der Kalkschwämme, 3 vols. (Berlin,
1872b), I, 484; idem, Natürliche Schöpfungsgeschichte, 3rd
ed. (Berlin, 1872a), p. xxxv. W. Harvey, “An Anatomical
Disquisition on the Motion of the Heart and Blood in
Animals,” in The Works of William Harvey, M.D., ed. and
trans. R. Willis (London, 1847), p. 82. The original essay
is in Latin. G. Heberer, ed., Der gerechtfertigte Haeckel
(Stuttgart, 1968). This is the most comprehensive modern
work available summarizing Haeckel's thought; it has ex-
tensive bibliographies. While it discusses the whole body
of his work, it devotes considerable space to recapitulation.
The major portion of the volume consists of long extracts
from Haeckel's own writings. W. His, Unsere Körperform
und das physiologische Problem ihrer Entstehung (Leipzig,
1874), pp. 168-71. C. J. Jung, Psychology of the Unconscious,
trans. B. M. Hinkle (New York, 1927), pp. 27-28. J. H. F.
Kohlbrugge, “Das biogenetische Grundgesetz. Eine
historische Studie,” Zoologischer Anzeiger, 37 (1911),
447-53. J. F. Meckel, System der vergleichenden Anatomie,
6 vols. (Halle, 1821-33), I, 396-97. W. Preyer, Die Seele
des Kindes (Leipzig, 1884), pp. vi-vii, 260-61. H. Spemann,
Forschung und Leben (Stuttgart, 1943). B. Spock, Baby and
Child Care, rev. ed. (New York, 1968), p. 229. E. Witschi,
Development of Vertebrates (Philadelphia, 1956), p. 6.

JANE OPPENHEIMER

[See also Biological Conceptions in Antiquity; Chain of
Being; Evolutionism; Inheritance of Acquired Characteris-
tics; Macrocosm and Microcosm.]