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In the
history of biology The history of biology traces the study of the living world from ancient to modern times. Although the concept of ''biology'' as a single coherent field arose in the 19th century, the biological sciences emerged from traditions of medicine an ...
, preformationism (or preformism) is a formerly popular theory that
organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells ( cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and fu ...
s develop from miniature versions of themselves. Instead of assembly from parts, preformationists believed that the
form Form is the shape, visual appearance, or configuration of an object. In a wider sense, the form is the way something happens. Form also refers to: *Form (document), a document (printed or electronic) with spaces in which to write or enter data * ...
of living things exist, in real terms, prior to their development.Maienschein, Jane,
Epigenesis and Preformationism
, The Stanford Encyclopedia of Philosophy (Fall 2008 Edition), Edward N. Zalta (ed.).
It suggests that all
organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells ( cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and fu ...
s were created at the same time, and that succeeding generations grow from homunculi, or
animalcule Animalcule ('little animal', from Latin ''animal'' + the diminutive suffix ''-culum'') is an old term for microscopic organisms that included bacteria, protozoans, and very small animals. The word was invented by 17th-century Dutch scientist ...
s, that have existed since the beginning of
creation Creation may refer to: Religion *''Creatio ex nihilo'', the concept that matter was created by God out of nothing *Creation myth, a religious story of the origin of the world and how people first came to inhabit it *Creationism, the belief that ...
, which is typically defined by religious beliefs. EpigenesisAccording to the
Oxford English Dictionary The ''Oxford English Dictionary'' (''OED'') is the first and foundational historical dictionary of the English language, published by Oxford University Press (OUP). It traces the historical development of the English language, providing a c ...
: It is also worth quoting this adumbration of the definition given there (viz., "The formation of an organic germ as a new product"):
(or neoformism), then, in this context, is the denial of preformationism: the idea that, in some sense, the form of living things comes into existence. As opposed to "strict" preformationism, it is the notion that "each embryo or organism is gradually produced from an undifferentiated mass by a series of steps and stages during which new parts are added" (Magner 2002, p. 154).Magner, Lois. ''A History of the Life Sciences''. New York: Marcel Dekker, Inc, 2002 This word is still used in a more modern sense, to refer to those aspects of the generation of form during ontogeny that are not strictly genetic, or, in other words, ''
epigenetic In biology, epigenetics is the study of stable phenotypic changes (known as ''marks'') that do not involve alterations in the DNA sequence. The Greek prefix '' epi-'' ( "over, outside of, around") in ''epigenetics'' implies features that are ...
''. Apart from those distinctions (preformationism-epigenesis and genetic-epigenetic), the terms ''preformistic development'', ''epigenetic development'' and '' somatic embryogenesis'' are also used in another context, in relation to the differentiation of a distinct
germ cell Germ or germs may refer to: Science * Germ (microorganism), an informal word for a pathogen * Germ cell, cell that gives rise to the gametes of an organism that reproduces sexually * Germ layer, a primary layer of cells that forms during embr ...
line. In preformistic development, the germ line is present since early development. In epigenetic development, the germ line is present, but it appears late. In somatic embryogenesis, a distinct germ line is lacking. Some authors call Weismannist development (either preformistic or epigenetic) that in which there is a distinct germ line. The historical ideas of preformationism and epigenesis, and the rivalry between them, are obviated by the contemporary understanding of the
genetic code The genetic code is the set of rules used by living cells to translate information encoded within genetic material ( DNA or RNA sequences of nucleotide triplets, or codons) into proteins. Translation is accomplished by the ribosome, which links ...
and its molecular basis together with
developmental biology Developmental biology is the study of the process by which animals and plants grow and develop. Developmental biology also encompasses the biology of regeneration, asexual reproduction, metamorphosis, and the growth and differentiation of st ...
and
epigenetics In biology, epigenetics is the study of stable phenotypic changes (known as ''marks'') that do not involve alterations in the DNA sequence. The Greek prefix '' epi-'' ( "over, outside of, around") in ''epigenetics'' implies features that are ...
.


Philosophical development

Pythagoras Pythagoras of Samos ( grc, Πυθαγόρας ὁ Σάμιος, Pythagóras ho Sámios, Pythagoras the Samian, or simply ; in Ionian Greek; ) was an ancient Ionian Greek philosopher and the eponymous founder of Pythagoreanism. His poli ...
is one of the earliest thinkers credited with ideas about the origin of form in the biological production of offspring. It is saide.g. by Ian Johnston, Malaspina University-College, Nanaimo, BC, in "...And Still We Evolve: A Handbook on the History of Modern Science"
Section Five: Heredity and Modern Genetics
, May 2000.
that he originated "spermism", the doctrine that fathers contribute the essential characteristics of their offspring while mothers contribute only a material substrate.
Aristotle Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of ...
accepted and elaborated this idea, and his writings are the vector that transmitted it to later Europeans. Aristotle purported to analyse ontogeny in terms of the material, formal, efficient, and teleological causes (as they are usually named by later anglophone philosophy) – a view that, though more complex than some subsequent ones, is essentially more epigenetic than preformationist. Later, European physicians such as
Galen Aelius Galenus or Claudius Galenus ( el, Κλαύδιος Γαληνός; September 129 – c. AD 216), often Anglicized as Galen () or Galen of Pergamon, was a Greek physician, surgeon and philosopher in the Roman Empire. Considered to be o ...
,
Realdo Colombo Matteo Realdo Colombo (c. 1515 – 1559) was an Italian professor of anatomy and a surgeon at the University of Padua between 1544 and 1559. Early life and education Matteo Realdo Colombo or Realdus Columbus, was born in Cremona, Lombardy, the ...
and
Girolamo Fabrici Girolamo Fabrici d'Acquapendente, also known as Girolamo Fabrizio or Hieronymus Fabricius (20 May 1533 – 21 May 1619), was a pioneering anatomist and surgeon known in medical science as "The Father of Embryology." Life and accomplishment ...
would build upon Aristotle's theories, which were prevalent well into the 17th century. In 1651,
William Harvey William Harvey (1 April 1578 – 3 June 1657) was an English physician who made influential contributions in anatomy and physiology. He was the first known physician to describe completely, and in detail, the systemic circulation and propert ...
published ''On the Generation of Animals'' ('' Exercitationes de Generatione Animalium''), a seminal work on
embryology Embryology (from Greek ἔμβρυον, ''embryon'', "the unborn, embryo"; and -λογία, '' -logia'') is the branch of animal biology that studies the prenatal development of gametes (sex cells), fertilization, and development of embr ...
that contradicted many of Aristotle's fundamental ideas on the matter. Harvey famously asserted, for example, that ''ex ovo omnia''—all animals come from eggs. Because of this assertion in particular, Harvey is often credited with being the father of ovist preformationism. However, Harvey's ideas about the process of development were fundamentally epigenesist.Clara Pinto Correia, ''The Ovary of Eve: Egg and Sperm Preformation'', Chicago: University of Chicago Press, 1997. As
gametes A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce ...
(male sperm and female ova) were too small to be seen under the best magnification at the time, Harvey's account of fertilization was theoretical rather than descriptive. Although he once postulated a "spiritous substance" that exerted its effect on the female body, he later rejected it as superfluous and thus unscientific. He guessed instead that fertilization occurred through a mysterious transference by contact, or contagion. Harvey's epigenesis, more mechanistic and less vitalist than the Aristotelian version, was, thus, more compatible with the
natural philosophy Natural philosophy or philosophy of nature (from Latin ''philosophia naturalis'') is the philosophical study of physics, that is, nature and the physical universe. It was dominant before the development of modern science. From the ancien ...
of the time. Still, the idea that unorganized matter could ultimately self-organize into life challenged the
mechanistic The mechanical philosophy is a form of natural philosophy which compares the universe to a large-scale mechanism (i.e. a machine). The mechanical philosophy is associated with the scientific revolution of early modern Europe. One of the first expo ...
framework of
Cartesianism Cartesianism is the philosophical and scientific system of René Descartes and its subsequent development by other seventeenth century thinkers, most notably François Poullain de la Barre, Nicolas Malebranche and Baruch Spinoza. Descartes is o ...
, which had become dominant in the
Scientific Revolution The Scientific Revolution was a series of events that marked the emergence of modern science during the early modern period, when developments in mathematics, physics, astronomy, biology (including human anatomy) and chemistry transforme ...
. Because of technological limitations, there was no available mechanical explanation for epigenesis. It was simpler and more convenient to postulate preformed miniature organisms that expanded in accordance with mechanical laws. So convincing was this explanation that some naturalists claimed to actually see miniature preformed animals (
animalcule Animalcule ('little animal', from Latin ''animal'' + the diminutive suffix ''-culum'') is an old term for microscopic organisms that included bacteria, protozoans, and very small animals. The word was invented by 17th-century Dutch scientist ...
s) in eggs and miniature plants in seeds. In the case of humans, the term
homunculus A homunculus ( , , ; "little person") is a representation of a small human being, originally depicted as small statues made out of clay. Popularized in sixteenth-century alchemy and nineteenth-century fiction, it has historically referred to the ...
was used.


Elaboration

After the discovery of
spermatozoa A spermatozoon (; also spelled spermatozoön; ; ) is a motile sperm cell, or moving form of the haploid cell that is the male gamete. A spermatozoon joins an ovum to form a zygote. (A zygote is a single cell, with a complete set of chromos ...
in 1677 by Dutch microscopist
Antonie van Leeuwenhoek Antonie Philips van Leeuwenhoek ( ; ; 24 October 1632 – 26 August 1723) was a Dutch microbiologist and microscopist in the Golden Age of Dutch science and technology. A largely self-taught man in science, he is commonly known as " the ...
, the epigenist theory proved more difficult to defend: How could complex organisms such as human beings develop from such simple organisms? Thereafter, Giuseppe degli Aromatari and then
Marcello Malpighi Marcello Malpighi (10 March 1628 – 30 November 1694) was an Italian biologist and physician, who is referred to as the "Founder of microscopical anatomy, histology & Father of physiology and embryology". Malpighi's name is borne by several ph ...
and
Jan Swammerdam Jan Swammerdam (February 12, 1637 – February 17, 1680) was a Dutch biologist and microscopist. His work on insects demonstrated that the various phases during the life of an insect— egg, larva, pupa, and adult—are different forms of the ...
made observations using microscopes in the late 17th century, and interpreted their findings to develop the preformationist theory. For two centuries, until the development of
cell theory In biology, cell theory is a scientific theory first formulated in the mid-nineteenth century, that living organisms are made up of cells, that they are the basic structural/organizational unit of all organisms, and that all cells come from pre ...
, preformationists would oppose epigenicists, and, inside the preformationist camp, spermists (who claimed the homunculus must come from the man) to ovists, who located the homunculus in the ova. Dutch microscopist
Antonie van Leeuwenhoek Antonie Philips van Leeuwenhoek ( ; ; 24 October 1632 – 26 August 1723) was a Dutch microbiologist and microscopist in the Golden Age of Dutch science and technology. A largely self-taught man in science, he is commonly known as " the ...
was one of the first to observe spermatozoa. He described the spermatozoa of about 30 species, and thought he saw in semen "all manner of great and small vessels, so various and so numerous that I do not doubt that they be nerves, arteries and veins...And when I saw them, I felt convinced that, in no full grown body, are there any vessels which may not be found likewise in semen." (Friedman 76-7)Friedman, David M. A Mind of Its Own: A Cultural History of the Penis. New York: The Free Press, 2001 Leeuwenhoek discovered that the origin of semen was the testicles and was a committed preformationist and spermist. He reasoned that the movement of spermatozoa was evidence of animal life, which presumed a complex structure and, for human sperm, a soul. (Friedman 79) In 1694, Nicolaas Hartsoeker, in his ''Essai de Dioptrique'' concerning things large and small that could be seen with optical lenses, produced an image of a tiny human form curled up inside the sperm, which he referred to in the French as ''petit l'infant'' and ''le petit animal''. This image, depicting what historians now refer to as the
homunculus A homunculus ( , , ; "little person") is a representation of a small human being, originally depicted as small statues made out of clay. Popularized in sixteenth-century alchemy and nineteenth-century fiction, it has historically referred to the ...
, has become iconic of the theory of preformationism, and appears in almost every textbook concerning the history of embryological science. Philosopher
Nicolas Malebranche Nicolas Malebranche ( , ; 6 August 1638 – 13 October 1715) was a French Oratorian Catholic priest and rationalist philosopher. In his works, he sought to synthesize the thought of St. Augustine and Descartes, in order to demonstrate the a ...
was the first to advance the hypothesis that each embryo could contain even smaller embryos
ad infinitum ''Ad infinitum'' is a Latin phrase meaning "to infinity" or "forevermore". Description In context, it usually means "continue forever, without limit" and this can be used to describe a non-terminating process, a non-terminating ''repeating'' pr ...
, like a Matryoshka doll. According to Malebranche, "an infinite series of plants and animals were contained within the seed or the egg, but only naturalists with sufficient skill and experience could detect their presence." (Magner 158-9) In fact, Malebranche only alleged this, observing that if microscopes enabled us to see very little animals and plants, maybe even smaller creatures could exist. He claimed that it was not unreasonable to believe that "they are infinite trees in only one seed," as he stated that we could already see chickens in eggs, tulips in bulbs, frogs in eggs. From this, he hypothesized that "all the bodies of humans and animals," already born and yet to be born, "were perhaps produced as soon as the creation of the world." Ova were known in some non-mammalian species, and
semen Semen, also known as seminal fluid, is an organic bodily fluid created to contain spermatozoa. It is secreted by the gonads (sexual glands) and other sexual organs of male or hermaphroditic animals and can fertilize the female ovum. Sem ...
was thought to spur the development of the preformed organism contained therein. The theory that located the homonculus in the egg was called ovism. But, when spermatozoa were discovered, a rival camp of spermists sprang up, claiming that the homunculus must come from the male. In fact, the term "spermatozoon," coined by
Karl Ernst von Baer Karl Ernst Ritter von Baer Edler von Huthorn ( – ) was a Baltic German scientist and explorer. Baer was a naturalist, biologist, geologist, meteorologist, geographer, and is considered a, or the, founding father of embryology. He was ...
, means "seed animals." With the discovery of sperm and the concept of spermism came a religious quandary. Why would so many little animals be wasted with each ejaculation of semen? Pierre Lyonet said the wastage proved that sperm could not be the seeds of life.
Leibniz Gottfried Wilhelm (von) Leibniz . ( – 14 November 1716) was a German polymath active as a mathematician, philosopher, scientist and diplomat. He is one of the most prominent figures in both the history of philosophy and the history of ma ...
supported a theory called
panspermism Panspermia () is the hypothesis, first proposed in the 5th century BCE by the Greek philosopher Anaxagoras, that life exists throughout the Universe, distributed by space dust, meteoroids, asteroids, comets, and planetoids, as well as by spacecraf ...
that the wasted sperm might actually be scattered (for example, by the wind) and generate life wherever they found a suitable host. Leibniz also believed that “death is only a transformation enveloped through diminution,” meaning that not only have organisms always existed in their living form, but that they will always exist, body united to soul, even past apparent death. In the 18th century, some animalculists thought that an animal's sperm behaved like the adult animal, and recorded such observations. Some, but not all, preformationists at this time claimed to see miniature organisms inside the sex cells. But, about this time, spermists began to use more abstract arguments to support their theories.
Jean Astruc Jean Astruc (19 March 1684, in Sauve, France – 5 May 1766, in Paris) was a professor of medicine in France at Montpellier and Paris, who wrote the first great treatise on syphilis and venereal diseases, and also, with a small anonymously publ ...
, noting that parents of both sexes seemed to influence the characteristics of their offspring, suggested that the animalcule came from the sperm and was then shaped as it passed into the egg. Buffon and Pierre Louis Moreau also advocated theories to explain this phenomenon. Preformationism, especially ovism, was the dominant theory of generation during the 18th century. It competed with
spontaneous generation Spontaneous generation is a superseded scientific theory that held that living creatures could arise from nonliving matter and that such processes were commonplace and regular. It was hypothesized that certain forms, such as fleas, could arise f ...
and epigenesis, but those two theories were often rejected on the grounds that inert matter could not produce life without God's intervention. Some animals' regenerative capabilities challenged preformationism, and Abraham Trembley's studies of the hydra convinced various authorities to reject their former views.
Lazaro Spallanzani Lazzaro Spallanzani (; 12 January 1729 – 11 February 1799) was an Italian Catholic priest (for which he was nicknamed Abbé Spallanzani), biologist and physiologist who made important contributions to the experimental study of bodily function ...
, Trembley's nephew, experimented with regeneration and semen, but failed to discern the importance of spermatozoa, dismissing them as parasitic worms and concluding instead that it was the liquid portion of semen that caused the preformed organism in the
ovum The egg cell, or ovum (plural ova), is the female reproductive cell, or gamete, in most anisogamous organisms (organisms that reproduce sexually with a larger, female gamete and a smaller, male one). The term is used when the female gamete i ...
to develop.


Criticisms and cell theory

Caspar Friedrich Wolff Caspar Friedrich Wolff (18 January 1733 – 22 February 1794) was a German physiologist and one of the founders of embryology. Life Wolff was born in Berlin, Brandenburg. In 1759 he graduated as an M.D. from the University of Halle with his dis ...
, an epigeneticist, was an 18th-century exception who argued for objectivity and freedom from religious influence on scientific questions. Despite careful observation of developing embryos, epigenesis suffered from a lack of a theoretical mechanism of generation. Wolff proposed an "essential force" as the agent of change, and
Immanuel Kant Immanuel Kant (, , ; 22 April 1724 – 12 February 1804) was a German philosopher and one of the central Enlightenment thinkers. Born in Königsberg, Kant's comprehensive and systematic works in epistemology, metaphysics, ethics, and ...
with Johann Friedrich Blumenbach proposed a "developing drive" or '' Bildungstrieb'', a concept related to
self-organization Self-organization, also called spontaneous order in the social sciences, is a process where some form of overall order arises from local interactions between parts of an initially disordered system. The process can be spontaneous when suffic ...
. Naturalists of the late 18th century and the 19th century embraced Wolff's philosophy, but primarily because they rejected the application of mechanistic development, as seen in the expansion of miniature organisms. It was not until the late 19th century that preformationism was discarded in the face of
cell theory In biology, cell theory is a scientific theory first formulated in the mid-nineteenth century, that living organisms are made up of cells, that they are the basic structural/organizational unit of all organisms, and that all cells come from pre ...
. Now, scientists "realized that they need not treat living organisms as machines, nor give up all hope of ever explaining the mechanisms that govern living beings." (Magner 173) When
John Dalton John Dalton (; 5 or 6 September 1766 – 27 July 1844) was an English chemist, physicist and meteorologist. He is best known for introducing the atomic theory into chemistry, and for his research into Color blindness, colour blindness, which ...
's
atomic theory Atomic theory is the scientific theory that matter is composed of particles called atoms. Atomic theory traces its origins to an ancient philosophical tradition known as atomism. According to this idea, if one were to take a lump of matter ...
of matter superseded Descartes' philosophy of
infinite divisibility Infinite divisibility arises in different ways in philosophy, physics, economics, order theory (a branch of mathematics), and probability theory (also a branch of mathematics). One may speak of infinite divisibility, or the lack thereof, of matter, ...
at the beginning of the 19th century, preformationism was struck a further blow. There was not enough space at the bottom of the spectrum to accommodate infinitely stacked animalcules, without bumping into the constituent parts of matter. (Gee 43) Gee, Henry. ''Jacob's Ladder: The History of the Human Genome''. New York: W.W. Norton & Company, Inc, 2004


Roux and Driesch

Near the end of the 19th century, the most prominent advocates of preformationatism and epigenesis were
Wilhelm Roux Wilhelm Roux (9 June 1850 – 15 September 1924) was a German zoologist and pioneer of experimental embryology. Early life Roux was born and educated in Jena, Germany where he attended university and studied under Ernst Haeckel. He also attende ...
and
Hans Driesch Hans Adolf Eduard Driesch (28 October 1867 – 17 April 1941) was a German biologist and philosopher from Bad Kreuznach. He is most noted for his early experimental work in embryology and for his neo-vitalist philosophy of entelechy. He has also ...
. Driesch's experiments on the development of the embryos of
sea urchin Sea urchins () are spiny, globular echinoderms in the class Echinoidea. About 950 species of sea urchin live on the seabed of every ocean and inhabit every depth zone from the intertidal seashore down to . The spherical, hard shells (tests) o ...
s are considered to have been decisively in favor of epigenesis.


See also

*
Biogenesis Spontaneous generation is a superseded scientific theory that held that living creatures could arise from nonliving matter and that such processes were commonplace and regular. It was hypothesized that certain forms, such as fleas, could arise ...


References


Bibliography

*
Edward Dolnick Edward Ishmael Dolnick (born November 10, 1952) is an American writer, formerly a science writer at the ''Boston Globe''. He has been published in ''Atlantic Monthly'', ''The New York Times Magazine'', and ''The Washington Post'', among other publ ...
, ''The Seeds of Life: From Aristotle to da Vinci, from Sharks' Teeth to Frogs' Pants, te Long and Strange Quest to Discover Where Babies Come From'', New York: Basic Books,2017, * Elizabeth B. Gasking, ''Investigations Into Generation 1651-1828'', Baltimore: The Johns Hopkins University Press, 1966 * Shirley A. Roe, ''Biology, Atheism, and Politics in 18th-century France,'' Chapter 2 pp. 36–60, in Alexander & Numbers, 2010 * Denis R. Alexander and Ronald L. Numbers (Eds), ''Biology and Ideology from Descartes to Dawkins,'' Chicago: University of Chicago Press, 2010, {{ISBN, 978-0-226-60840-2 History of biology Philosophy of science Obsolete biology theories