Preformationism
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In the history of biology, preformationism (or preformism) is a formerly popular theory that
organism An organism is any life, living thing that functions as an individual. Such a definition raises more problems than it solves, not least because the concept of an individual is also difficult. Many criteria, few of them widely accepted, have be ...
s develop from miniature versions of themselves. Instead of assembly from parts, preformationists believed that the form 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.).
Preformationists suggested that all
organism An organism is any life, living thing that functions as an individual. Such a definition raises more problems than it solves, not least because the concept of an individual is also difficult. Many criteria, few of them widely accepted, have be ...
s were created at the same time, and that succeeding generations grow from homunculi, or animalcules, that have existed since the beginning of creation, which is typically defined by religious beliefs. EpigenesisAccording to the
Oxford English Dictionary The ''Oxford English Dictionary'' (''OED'') is the principal historical dictionary of the English language, published by Oxford University Press (OUP), a University of Oxford publishing house. The dictionary, which published its first editio ...
: 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''. 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 A germ cell is any cell that gives rise to the gametes of an organism that reproduces sexually. In many animals, the germ cells originate in the primitive streak and migrate via the gut of an embryo to the developing gonads. There, they unde ...
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 Genetic code is a set of rules used by living cell (biology), cells to Translation (biology), translate information encoded within genetic material (DNA or RNA sequences of nucleotide triplets or codons) into proteins. Translation is accomplished ...
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 (biology), regeneration, asexual reproduction, metamorphosis, and the growth and di ...
and
epigenetics In biology, epigenetics is the study of changes in gene expression that happen without changes to the DNA sequence. The Greek prefix ''epi-'' (ἐπι- "over, outside of, around") in ''epigenetics'' implies features that are "on top of" or "in ...
.


Philosophical development

Pythagoras Pythagoras of Samos (;  BC) was an ancient Ionian Greek philosopher, polymath, and the eponymous founder of Pythagoreanism. His political and religious teachings were well known in Magna Graecia and influenced the philosophies of P ...
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 (; 384–322 BC) was an Ancient Greek philosophy, Ancient Greek philosopher and polymath. His writings cover a broad range of subjects spanning the natural sciences, philosophy, linguistics, economics, politics, psychology, a ...
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 (; September 129 – AD), often Anglicization, anglicized as Galen () or Galen of Pergamon, was a Ancient Rome, Roman and Greeks, Greek physician, surgeon, and Philosophy, philosopher. Considered to be one o ...
, Realdo Colombo and Girolamo Fabrici 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 to anatomy and physiology. He was the first known physician to describe completely, and in detail, pulmonary and systemic circulation ...
published ''On the Generation of Animals'' ('' Exercitationes de Generatione Animalium''), a seminal work on
embryology Embryology (from Ancient Greek, Greek ἔμβρυον, ''embryon'', "the unborn, embryo"; and -λογία, ''-logy, -logia'') is the branch of animal biology that studies the Prenatal development (biology), prenatal development of gametes (sex ...
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 (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, while ignoring any supernatural influence. It was dominant before the develop ...
of the time. Still, the idea that unorganized matter could ultimately self-organize into life challenged the mechanistic 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 i ...
, which had become dominant in the
Scientific Revolution The Scientific Revolution was a series of events that marked the emergence of History of science, modern science during the early modern period, when developments in History of mathematics#Mathematics during the Scientific Revolution, mathemati ...
. 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 ( animalcules) in eggs and miniature plants in seeds. In the case of humans, the term homunculus was used.


Elaboration

After the discovery of
spermatozoa A spermatozoon (; also spelled spermatozoön; : spermatozoa; ) is a motile sperm cell (biology), cell produced by male animals relying on internal fertilization. A spermatozoon is a moving form of the ploidy, haploid cell (biology), cell that is ...
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 art, science and technology. A largely self-taught man in science, he is commonly known as " ...
, 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 Giuseppe degli Aromatari or Josephus de Aromatariis (March 25, 1587 – July 16, 1660) was an Italian physician and writer. Biography Giuseppe degli Aromatari was born to Favorino and Filogenia Paolucci, from a noble and ancient family at Assi ...
and then
Marcello Malpighi Marcello Malpighi (10 March 1628 – 30 November 1694) was an Italians, Italian biologist and physician, who is referred to as the "founder of microscopical anatomy, histology and father of physiology and embryology". Malpighi's name is borne by ...
and Jan Swammerdam 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 pr ...
, 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 art, science and technology. A largely self-taught man in science, he is commonly known as " ...
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, has become iconic of the theory of preformationism, and appears in almost every textbook concerning the history of embryological science. Philosopher Nicolas Malebranche was the first to advance the hypothesis that each embryo could contain even smaller embryos ad infinitum, 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 a bodily fluid that contains spermatozoon, spermatozoa which is secreted by the male gonads (sexual glands) and other sexual organs of male or hermaphrodite, hermaphroditic animals. In humans and placen ...
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, 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 supported a theory called panspermism 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 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, 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 to develop.


Criticisms and cell theory

Caspar Friedrich Wolff, 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 (born Emanuel Kant; 22 April 1724 – 12 February 1804) was a German Philosophy, philosopher and one of the central Age of Enlightenment, Enlightenment thinkers. Born in Königsberg, Kant's comprehensive and systematic works ...
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 and disorder, order arises from local interactions between parts of an initially disordered system. The process can be spont ...
. 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 pr ...
. 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 introduced the atomic theory into chemistry. He also researched Color blindness, colour blindness; as a result, the umbrella term ...
's
atomic theory Atomic theory is the scientific theory that matter is composed of particles called atoms. The definition of the word "atom" has changed over the years in response to scientific discoveries. Initially, it referred to a hypothetical concept of ...
of matter superseded Descartes' philosophy of infinite divisibility 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 and Hans Driesch. Driesch's experiments on the development of the embryos of
sea urchin Sea urchins or urchins () are echinoderms in the class (biology), class Echinoidea. About 950 species live on the seabed, inhabiting all oceans and depth zones from the intertidal zone to deep seas of . They typically have a globular body cove ...
s are considered to have been decisively in favor of epigenesis.


See also

*
Biogenesis Spontaneous generation is a Superseded scientific theories, superseded scientific theory that held that living creatures could arise from abiotic component, non-living matter and that such processes were commonplace and regular. It was Hypoth ...


References


Bibliography

* Edward Dolnick, ''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