August Weismann

For the 19th-century New York politician, see Augustus Weismann.

August Friedrich Leopold Weismann (17 January 1834 – 5 November 1914) was
August Weismann
a German evolutionary biologist. Ernst Mayr ranked him as the second most notable
evolutionary theorist of the 19th century, after Charles Darwin. Weismann became
the Director of the Zoological Institute and the first Professor of Zoology at
Freiburg.

His main contribution involved germ plasm theory, at one time also known as
Weismannism,[1] according to which inheritance (in a multicellular organism) only
takes place by means of the germ cells—the gametes such as egg cells and sperm
cells. Other cells of the body—somatic cells—do not function as agents of heredity.
The effect is one-way: germ cells produce somatic cells and are not affected by
anything the somatic cells learn or therefore any ability an individual acquires during
its life. Genetic information cannot pass from soma to germ plasm and on to the next
generation. Biologists refer to this concept as the Weismann barrier.[2] This idea, if
true, rules out the inheritance of acquired characteristics as proposed by Jean-
Baptiste Lamarck.[3]

The idea of the Weismann barrier is central to the modern synthesis of the early 20th Born 17 January 1834
century, though scholars do not express it today in the same terms. In Weismann's Frankfurt am Main
opinion the largely random process of mutation, which must occur in thegametes (or Died 5 November 1914
stem cells that make them) is the only source of change for natural selection to work (aged 80)
on. Weismann became one of the first biologists to deny Lamarckism entirely.[4] Freiburg
Weismann's ideas preceded the rediscovery of Gregor Mendel's work, and though
Known for germ plasm theory
Weismann was cagey about accepting Mendelism, younger workers soon made the
connection.
Awards Darwin–Wallace Medal
(Silver, 1908)
Weismann is much admired today. Ernst Mayr judged him to be the most important
evolutionary thinker between Darwin and the evolutionary synthesis around 1930–1940, and "one of the great biologists of all
time".[5]

Contents
Life
Youth and studies
Professional life
Contributions to evolutionary biology
German work on cells
1868–1881/82
1882–1895
1896–1910
Experiments on the inheritance of mutilations
Awards
Publications by Weismann
 References
Sources
External links

Life

Youth and studies

Weismann was born a son of high school teacher Johann (Jean) Konrad Weismann (1804–1880), a graduate of ancient languages and
theology, and his wife Elise (1803–1850), née Lübbren, the daughter of the county councillor and mayor of Stade, on 17 January
1834 in Frankfurt am Main. He had a typical 19th century bourgeois education, receiving music lessons from the age of four, and
drafting and painting lessons from Jakob Becker (1810–1872) at the Frankfurter Städelsche Institut from the age of 14. His piano
teacher was a devoted butterfly collector and introduced him to the collecting of imagos and caterpillars. But studying natural
sciences was out of the question due to the cost involved and limited job prospects. A friend of the family, chemist Friedrich Wöhler
(1800–1882), recommended studying medicine. A foundation from the inheritance of Weismann's mother allowed him to take up
studies in Göttingen. Following his graduation in 1856, he wrote his dissertation on the synthesis of
hippuric acid in the human body.

Professional life
Immediately after university, Weismann took on a post as assistant at the Städtische Klinik (city clinic) in Rostock. Weismann
successfully submitted two manuscripts, one about hippuric acid in herbivores, and one about the salt content of the Baltic Sea, and
won two prizes. The paper about the salt content dissuaded him from becoming a chemist, since he felt himself lacking in
apothecarial accuracy.

After a study visit to see Vienna's museums and clinics, he graduated as a physician and settled in Frankfurt with a medical practice
in 1868. During the war between Austria, France and Italy in 1859, he became Chief Medical Officer in the military. During a leave
from duty, he walked through Northern Italy and the County of Tyrol. After a sabbatical in Paris, he worked with Rudolf Leuckart at
the University of Gießen. He returned to Frankfurt as personal physician to the banishedArchduke Stephen of Austriaat Schaumburg
Castle from 1861 to 1863.

From 1863, he was privatdozent in comparative anatomy and zoology; from 1866 extraordinary professor; and from 1873 to 1912
full professor, first holder of the chair in zoology and director of the zoological institute at Albert Ludwig University of Freiburg in
Breisgau. He retired in 1912.[6]

His son Julius Weismann (1879-1950) was a composer.

Contributions to evolutionary biology

At the beginning of Weismann's preoccupation with evolutionary theory was his grappling with Christian creationism as a possible
alternative. In his workÜber die Berechtigung der Darwin'schen Theorie (On the justification of the Darwinian theory) he compared
creationism and evolutionary theory, and concluded that many biological facts can be seamlessly accommodated within evolutionary
theory, but remain puzzling if considered the result of acts of creation.

After this work, Weismann accepted evolution as a fact on a par with the fundamental assumptions of astronomy (e.g.
Heliocentrism).
Weismann's position towards the mechanism of inheritance and its role for evolution changed during his life. Three periods can be
distinguished.

German work on cells

Weismann's work on the demarcation between germ-line and
soma can scarcely be appreciated without considering the work
of (mostly) German biologists during the second half of the
19th century. This was the time that the mechanisms of cell
division began to be understood. Eduard Strasburger, Walther
Flemming, Heinrich von Waldeyer and the Belgian Edouard
Van Beneden laid the basis for the cytology and cytogenetics of
the 20th century. Strasburger, the outstanding botanical
physiologist of that century, coined the terms nucleoplasm and
cytoplasm. He said "new cell nuclei can only arise from the Weismann's germ plasm theory. The hereditary material,
the germ plasm, is confined to thegonads. Somatic cells
division of other cell nuclei". Van Beneden discovered how
(of the body) develop afresh in each generation from the
chromosomes combined at meiosis, during the production of germ plasm.
gametes, and discovered and named chromatin. Walther
Flemming, the founder of cytogenetics, named mitosis, and
pronounced "omnis nucleus e nucleo" (which means the same as Strasburger's dictum). The discovery of mitosis, meiosis and
chromosomes is regarded as one of the 100 most important scientific discoveries of all times,[7] and one of the 10 most important
discoveries in cell biology.[8]

Meiosis was discovered and described for the first time in sea urchin eggs in 1876, by Oscar Hertwig. It was described again in 1883,
at the level of chromosomes, by Van Beneden in Ascaris eggs. The significance of meiosis for reproduction and inheritance,
however, was first described in 1890 by Weismann, who noted that two cell divisions were necessary to transform one diploid cell
into four haploid cells if the number of chromosomes had to be maintained. Thus the work of the earlier cytologists laid the ground
for Weismann, who turned his mind to the consequences for evolution, which was an aspect the cytologists had not addressed.[9] All
this took place before the work of Mendel had been rediscovered

1868–1881/82
Weismann starts out believing, like many other 19th century scientists, among them Charles Darwin, that the observed variability of
individuals of one species is due to the inheritance of sports (Darwin's term). He believed, as written in 1876, that transmutation of
species is directly due to the influence of environment. He also wrote, "if every variation is regarded as a reaction of the organism to
external conditions, as a deviation of the inherited line of development, it follows that no evolution can occur without a change of the
environment". (this is close to the modern use of the concept that changes in the environment can mediate selective pressures on a
population, so leading to evolutionary change.) W
eismann also used the classicLamarckian metaphor of use and disuse of an organ.

1882–1895
Weismann's first rejection of the inheritance of acquired traits was in a lecture in 1883, titled "On inheritance" ("Über die
Vererbung"). Again, as in his treatise on creation vs. evolution, he attempts to explain individual examples with either theory. For
instance, the existence of non-reproductive castes of ants, such as workers and soldiers, cannot be explained by inheritance of
acquired characters. Germ plasm theory, on the other hand, does so effortlessly. Weismann used this theory to explain Lamark's
original examples for "use and disuse", such as the tendency to have degenerate wings and stronger feet in domesticated waterfowl.

1896–1910
Weismann worked on the embryology of sea urchin eggs, and in the course of this observed different kinds of cell division, namely
equatorial division and reductional division, terms he coinedÄquatorialteilung
( and Reduktionsteilung respectively).

His germ plasm theory states that multicellular organisms consist of germ cells containing heritable information, and somatic cells
that carry out ordinary bodily functions. The germ cells are influenced neither by environmental influences nor by learning or
morphological changes that happen during the lifetime of an organism, which information is lost after each generation. The concept
as he proposed it was referred to as Weismannism in his day, for example in the book An examination of Weismannism by George
Romanes[1] This idea was illuminated and explained by the rediscovery of Gregor Mendel's work in the early years of the 20th
century (see Mendelian inheritance).

Experiments on the inheritance of mutilations

The idea that germline cells contain information that passes to each generation unaffected by experience and independent of the
somatic (body) cells, came to be referred to as the Weismann barrier, and is frequently quoted as putting a final end to the theory of
Lamarck and the inheritance of acquired characteristics. What Lamarck claimed was the inheritance of characteristics acquired
through effort, or will.

Weismann conducted the experiment of removing the tails of 68 white mice, repeatedly over 5 generations, and reporting that no
mice were born in consequence without a tail or even with a shorter tail. He stated that "901 young were produced by five generations
of artificially mutilated parents, and yet there was not a single example of a rudimentary tail or of any other abnormality in this
organ."[10] Weismann was aware of the limitations of this experiment, and made it clear that he embarked on the experiment
precisely because, at the time, there were many claims of animals inheriting mutilations (he refers to a claim regarding a cat that had
lost its tail having numerous tail-less offspring). There were also claims of Jews born without foreskins. None of these claims, he
said, were backed up by reliable evidence that the parent had in fact been mutilated, leaving the perfectly plausible possibility that the
modified offspring were the result of a mutated gene. The purpose of his experiment was to lay the claims of inherited mutilation to
rest. The results were consistent with Weismann's germ plasm theory.

Awards
He was awarded the Linnean Society of London's Darwin-Wallace Medal in 1908.

Publications by Weismann
1868. Über die Berechtigung der Darwin'schen Theorie : Ein akademischer Vortrag gehalten am 8. Juli 1868 in der
Aula der Universität zu Freiburg im Breisgau. Engelmann, Leipzig.
1872. Über den Einfluß der Isolierung auf die Artbildung
. Engelmann, Leipzig.
1875. Studien zur Descendenz-Theorie. I. Ueber den Saison-Dimorphismus der Schmetterlinge . Leipzig.
1876. Studien zur Descendenztheorie: II. Ueber die letzten Ursachen der ransmutationen.
T Leipzig.
1883. Die Entstehung der Sexualzellen bei den Hydromedusen: Zugleich ein Beitrag zur Kenntniss des Baues und
der Lebenserscheinungen dieser Gruppe. Fischer, Jena.
1885. Die Continuität des Keimplasmas als Grundlage einer Theorie der ererbung.
V Fischer, Jena.
1887. Zur Frage nach der Vererbung erworbener Eigenschaften. In:Biol. Zbl. 6:33–48
1887. Über die Zahl der Richtungskörper und über ihre Bedeutung für die ererbung.
V Fischer, Jena.
1889 Translations: Essays upon Heredity Oxford Clarendon Press – Full online text[11]
1892. Das Keimplasma: eine Theorie der Vererbung. Fischer, Jena.
1893 Translation: Germ-Plasm, a theory of HeredityCharles Scribner's Sons – Full online text
1892. Aufsätze über Vererbung und angewandet biologische Fragen. Fischer, Jena.
1893. Die Allmacht der Naturzüchtung: eine Erwiderung anHerbert Spencer. Jena.
1902. Vorträge über Deszendenztheorie: Gehalten an der Universität zu Freiburg im Breisgau. Fischer
, Jena. 2
volums

References
1. Romanes, George John.An examination of Weismannism. The Open court publishing company in Chicago 1893[1]
(https://openlibrary.org/books/OL23380098M/An_examination_of_Weismannism)
2. Germ-Plasm, a theory of heredity (1893)- Full online text(http://www.esp.org/books/weismann/germ-plasm/facsimil
e/). Esp.org. Retrieved on 2012-02-25.
3. Huxley, Julian 1942. Evolution, the modern synthesis. p. 17
 4. Essays upon heredity (1889) Oxford Clarendon Press – Full online text(http://www.esp.org/books/weismann/essays/
facsimile/). Esp.org. Retrieved on 2012-02-25.
5. Mayr, Ernst 1982. The growth of biological thought. Harvard. p. 698
6. Gaup, Ernst 1917. August Weismann: sein Leben und sein Werk. Fischer, Jena.
7. 100 Greatest Discoveries – Carnegie Institution(http://carnegieinstitution.org/cover/top_100/)Archived (https://web.a
rchive.org/web/20070927070422/http://carnegieinstitution.org/cover/top_100/) 2007-09-27 at the Wayback Machine.
at carnegieinstitution.org
8. The Science Channel :: 100 Greatest Discoveries: Biology(http://science.discovery.com/convergence/100discoverie
s/big100/biology.html) Archived (https://web.archive.org/web/20061024155730/http://science.discovery
.com/converg
ence/100discoveries/big100/biology.html) 2006-10-24 at the Wayback Machine. at science.discovery.com
9. Although, of course, Ernst Haeckel had; but he was not a cytologist.
10. Tollefsbol, Trygve (2017). Handbook of Epigenetics: The New Molecular and Medical Genetics(https://books.google.
com/books?id=uJupDQAAQBAJ&pg=P A234). Elsevier Science. p. 234.ISBN 978-0-12-805477-2. Originally
published in Weismann's 1889 Essays Upon Heredity (http://www.esp.org/books/weismann/essays/facsimile/).
11. "Review of Essays upon Heredity and kindred Biological Problemsby August Weismann" (https://babel.hathitrust.or
g/cgi/pt?id=coo.31924065563631;view=1up;seq=382) . The Quarterly Review. 170: 370–393. April 1890.

Sources
Churchill F.B. 1968. August Weismann and a break from tradition.J. Hist. Biol. 1, 91–112.
Churchill F.B. 1970. Hertwig, Weismann, and the meaning of the reduction division, circa 1890.Isis 61, 429–457.
Löther, Rolf 1990. Wegbereiter der Genetik: Gregor Johann Mendel und August Weismann. Verlag Harri Deutsch,
Frankfurt am Main. ISBN 3-8171-1130-4
Risler H. 1968. August Weismann 1834–1914. In: Berichte der Naturforschenden Gesellschaft Freiburg im Breisgau .
77–93
Risler H. 1985. August Weismanns Leben und Wirken nach Dokumenten aus seinem Nachlass.In: Freiburger
Universitätsblätter Heft 87/88, Freiburg. 23–42
Romanes, George John 1893. An Examination of W eismannism. London, Longmans.

External links
Media related to August Weismann at Wikimedia Commons
Works written by or aboutAugust Weismann at Wikisource
Works by August Weismann at Project Gutenberg
Works by or about August Weismann at Internet Archive
Biography, bibliography and access to digitalsources in the Virtual Laboratory of the Max Planck Institute for the
History of Science

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