Pangenesis: gemmules from the whole body gather in the germ cells

1868 Life and health theory theory refuted

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A hypothesis that every cell of the body throws off minute particles, gemmules, which circulate freely through the organism, multiply by division and gather in the reproductive products, where cells like their parents later develop from them.

The hypothesis was put forward in 1868 by the Englishman Charles Darwin, to connect 'under a single point of view' reproduction, regeneration, reversion, the action of the first male on the dam, the inheritance of mutilations and of the effects of use and disuse, that is, both telegony and Lamarck's law. Darwin himself called it provisional and granted that 'the existence of free gemmules is a gratuitous assumption'. The first test came from within the family: his cousin Galton transfused the blood of rabbits and found no gemmules.

Then the hypothesis was displaced by Weismann's germ plasm and, after 1900, by Mendelian heredity. The dead end left a trace: in the Dutchman Hugo de Vries's 'intracellular pangenesis' of 1889 and the word 'pangene', from which 'gene' grew.

Why this date

High for the year. The 'provisional hypothesis of pangenesis' forms chapter XXVII of the second volume of 'The Variation of Animals and Plants under Domestication' (London: John Murray, 1868), pp. 357-404 of the first edition; the statement of the hypothesis begins on p. 374. Darwin had shown a manuscript of the hypothesis to Huxley as early as 1865 (Holterhoff, 2014), but the axis takes the year of printing.

What it grew from

• Inheritance of acquired characters: the second law of the 'Philosophie zoologique'

  influenced · Confidence: medium

  Pangenesis was built so as to explain, among other things, the inheritance of the acquired: a cell altered by conditions or by use throws off altered gemmules. Lamarck's law entered the list of facts that Darwin's hypothesis undertook to connect.

  Dispute: The confidence is deliberately lowered. Historians agree that pangenesis explained the inheritance of acquired characters, but not that Darwin took this from Lamarck in particular: Burkhardt insists that for Lamarck and his contemporaries this was a commonplace rather than Lamarck's 'signature' idea, and that Darwin held it in his own right from the time of the Origin. Cowan traces the line through Buffon and Erasmus Darwin. The link runs from the doctrine first written down as a law in 1809 to the hypothesis that undertook to explain it, not from book to book.

  Link sources

  • Ruth Schwartz Cowan, Commentary: Before Weismann and germplasm there was Galton and eugenics, International Journal of Epidemiology 45 (1), 15-20 (2016) Verified 20 Sep 2026

    Quite explicitly, pangenesis allowed for and explained how acquired characters could be inherited.

  • Richard W. Burkhardt, Lamarck, Evolution, and the Inheritance of Acquired Characters, Genetics 194 (4), 793-805 (2013) Verified 20 Sep 2026

    Nine years later in his Variation of Animals and Plants Under Domestication, Darwin offered his “provisional hypothesis of pangenesis” to account for a whole range of phenomena related to heredity and development, including the inheritance of acquired characters (Darwin 1868).

  • Yongsheng Liu, Qi Chen, 150 years of Darwin's theory of intercellular flow of hereditary information, Nature Reviews Molecular Cell Biology 19 (12), 749-750 (2018) Verified 20 Sep 2026

    Importantly, Darwin proposed that the environment could modify gemmules, and thus that the composition of gemmules mirrors the organism’s environmental exposure.

  "Verified" means that on that day the quotation was found at the address given, word for word. No date means we have not been able to open the source yet, not that the quotation is wrong.

• Telegony: Lord Morton's mare and the 'infection' of the dam by her first mate

  influenced · Confidence: medium

  Darwin accepted Morton's case as 'perfectly well-ascertained' and included the action of the first male on the dam among the facts pangenesis had to explain: gemmules of the quagga lodged in the mare's ovaries went on marking her offspring by other stallions.

  Dispute: The main source here is Darwin himself rather than a historian: the words about the 'perfectly well-ascertained' case come from the chapter on pangenesis. Among historians, Burkhardt lists 'the influence of the male element on the female form' among the phenomena pangenesis was built to cover, and Ewart, who refuted telegony himself, testifies that Darwin held Morton's case to be strong evidence. No direct statement by a historian that pangenesis would have looked different without belief in telegony was found; the link is marked as influence, not as a prerequisite.

  Link sources

  • Charles Darwin, The Variation of Animals and Plants under Domestication, London: John Murray, 1868, vol. 2, ch. XXVII, p. 366 (Darwin Online, F877) (1868) Verified 20 Sep 2026

    But we have the analogous and perfectly well-ascertained case of the male element of a distinct form, as with the quagga and Lord Morton's mare, affecting the ovarium of the female, so that the ovules and offspring subsequently produced by her when impregnated by other males are plainly affected and hybridised by the first male.

  • Richard W. Burkhardt, Lamarck, Evolution, and the Inheritance of Acquired Characters, Genetics 194 (4), 793-805 (2013) Verified 20 Sep 2026

    In his “provisional hypothesis of pangenesis,” he postulated that small, material particles that he called “gemmules” were produced in all the cells of the body, subsequently collected in the gonads and transmitted in reproduction, and accounted for a remarkable battery of hereditary and developmental phenomena, including (among others) reversion, “prepotency,” “the influence of the male element on the female form,”

  • J. C. Ewart, «Telegony», Encyclopaedia Britannica, 11th ed., vol. 26 (1911) Verified 20 Sep 2026

    Darwin and others having regarded Lord Morton’s mare as affording very strong evidence in support of the infection hypothesis, it was considered some years ago desirable to repeat Lord Morton’s experiment as accurately as possible.

  "Verified" means that on that day the quotation was found at the address given, word for word. No date means we have not been able to open the source yet, not that the quotation is wrong.

Other links

• Blood transfusion in rabbits: no gemmules in the blood

  refuted by · Confidence: high

  Galton tested the only plausible route of gemmules through the body, the blood: after massive transfusions from rabbits of other breeds, 124 offspring in 21 litters came out pure-bred. He concluded that the blood carries no gemmules, and scientific opinion on pangenesis, historians say, turned quickly to scepticism thereafter.

  Dispute: The dispute took place at once and in print. Darwin replied in Nature on 27 April 1871 that in the chapter on pangenesis he had 'not said one word about the blood, or about any fluid proper to any circulating system', that the hypothesis covers Protozoa and plants, which have no blood, and that Galton's conclusion was 'a little hasty'; Cowan calls this nit-picking, since Darwin had written of 'the steady circulation of fluids throughout the body' and in a year of correspondence never questioned the design of the experiment. Galton apologised on 4 May 1871 and closed his letter with 'Vive Pangenesis'. Karl Pearson in 'The Grammar of Science' (2nd ed., 1900, p. 335) objected that pangenesis is no more disproved by gemmules not found in the blood than the atomic theory by atoms not found in the air (given after the English Wikipedia article 'Pangenesis'; the page of the book was not checked). What finally removed the hypothesis from the agenda was not the rabbits but Weismann's germ plasm and the rediscovery of Mendel in 1900; here Galton's experiment is taken as the first refutation and the germ plasm as the second.

  Link sources

  • Ruth Schwartz Cowan, Commentary: Before Weismann and germplasm there was Galton and eugenics, International Journal of Epidemiology 45 (1), 15-20 (2016) Verified 20 Sep 2026

    the results were negative; out of 124 offspring in 21 litters born by transfused rabbit does, not a single ‘mongrel’ had appeared. Galton presented a full report of his findings to the Royal Society in March, 1871.

  • Yongsheng Liu, Qi Chen, 150 years of Darwin's theory of intercellular flow of hereditary information, Nature Reviews Molecular Cell Biology 19 (12), 749-750 (2018) Verified 20 Sep 2026

    In a series of blood transfusion experiments, Francis Galton failed to detect fur colour changes in the offspring of silver-grey rabbits that were transfused with the blood of lop-eared rabbits; he concluded that the blood did not carry Darwin’s hypothetical gemmules.

  • Charles Darwin, Pangenesis, Nature 3 (27 April 1871), 502-503 (Darwin Online, F1751) (1871)

    These experiments were undertaken to test whether there was any truth in my provisional hypothesis of Pangenesis.

  "Verified" means that on that day the quotation was found at the address given, word for word. No date means we have not been able to open the source yet, not that the quotation is wrong.

• 'Das Keimplasma': the continuity of the germ plasm and the mouse-tail experiment

  refuted by · Confidence: medium

  Weismann's germ plasm cancelled the very premise of pangenesis, the flow of hereditary matter from the body to the germ cells: for Weismann heredity runs only from the germ plasm to the body. Weismann called gemmules 'a pure invention', and pangenesis, in the words of Liu and Chen, was quickly replaced by his theory.

  Dispute: The confidence is medium: the authors of the main quotation, Liu and Chen, are biologists rather than historians, and their piece argues for a 'resurrection' of pangenesis through extracellular vesicles and RNA; Robinson's DSB article speaks of the denial of inheritance of the acquired, not of pangenesis by name. Historians also differ on what exactly closed pangenesis: de Beer places the end at the rediscovery of Mendel in 1900, Julian Huxley at Morgan's chromosome theory. Moreover, pangenesis has living offspring: de Vries's 'intracellular pangenesis' (1889) kept the particles but gave up their transport from the body to the germ cells. It is that transport that is marked as the dead end, not the idea of hereditary particles.

  Link sources

  • Yongsheng Liu, Qi Chen, 150 years of Darwin's theory of intercellular flow of hereditary information, Nature Reviews Molecular Cell Biology 19 (12), 749-750 (2018) Verified 20 Sep 2026

    The Pangenesis theory was quickly replaced by Weismann’s Germ-Plasm theory, which postulated that information only flows from germ cells to somatic cells, and not vice versa (this is also known as the ‘Weismann barrier’).

  • Yongsheng Liu, Qi Chen, 150 years of Darwin's theory of intercellular flow of hereditary information, Nature Reviews Molecular Cell Biology 19 (12), 749-750 (2018) Verified 20 Sep 2026

    These speculations could not be validated at the time and were considered as “the fictions of the fancy” by the physician and microscopist Lionel Beale, and as “a pure invention” by the evolutionary biologist August Weismann.

  • Gloria Robinson, «Weismann, August Friedrich Leopold», Complete Dictionary of Scientific Biography (2008) Verified 5 Oct 2026

    He maintained that the germ plasm, the substance of heredity, was transmitted from generation to generation, distinguishing it from the somatoplasm; he also was foremost in his day in denying that acquired characters were inherited.

  "Verified" means that on that day the quotation was found at the address given, word for word. No date means we have not been able to open the source yet, not that the quotation is wrong.

Node sources

• Charles Darwin, The Variation of Animals and Plants under Domestication, London: John Murray, 1868, vol. 2, ch. XXVII, p. 374 (Darwin Online, F877) (1868) Verified 20 Sep 2026

  But besides this means of increase I assume that cells, before their conversion into completely passive or "formed material," throw off minute granules or atoms, which circulate freely throughout the system, and when supplied with proper nutriment multiply by self-division, subsequently becoming developed into cells like those from which they were derived. These granules for the sake of distinctness may be called cell-gemmules, or, as the cellular theory is not fully established, simply gemmules.

• Charles Darwin, The Variation of Animals and Plants under Domestication, London: John Murray, 1868, vol. 2, ch. XXVII, p. 357, (Darwin Online, F877) (1868) Verified 20 Sep 2026

  THE FACTS TO BE CONNECTED UNDER A SINGLE POINT OF VIEW, NAMELY, THE VARIOUS KINDS OF REPRODUCTION

• Richard W. Burkhardt, Lamarck, Evolution, and the Inheritance of Acquired Characters, Genetics 194 (4), 793-805 (2013) Verified 20 Sep 2026

  In his “provisional hypothesis of pangenesis,” he postulated that small, material particles that he called “gemmules” were produced in all the cells of the body, subsequently collected in the gonads and transmitted in reproduction, and accounted for a remarkable battery of hereditary and developmental phenomena, including (among others) reversion, “prepotency,” “the influence of the male element on the female form,” organs developed in the wrong place, the regeneration of lost parts, the direct action of changed conditions, and “the inherited effects of the use or disuse of particular organs.”

• Kate Holterhoff, The History and Reception of Charles Darwin's Hypothesis of Pangenesis, Journal of the History of Biology 47 (4), 661-695 (2014) Verified 20 Sep 2026

  First published in The Variation of Animals and Plants under Domestication (1868), pangenesis stated that inheritance can be explained by sub-cellular “gemmules” which aggregated in the sexual organs during intercourse.

• Yongsheng Liu, Qi Chen, 150 years of Darwin's theory of intercellular flow of hereditary information, Nature Reviews Molecular Cell Biology 19 (12), 749-750 (2018) Verified 20 Sep 2026

  Pangenesis provided a potential explanation for a range of phenomena pertaining to inheritance and phenotypic variation, including ‘prepotency’ (analogous to Gregor Mendel’s ‘dominance’, introduced in 1866), graft hybridization, reversion and the inheritance of acquired characteristics

"Verified" means that on that day the quotation was found at the address given, word for word. No date means we have not been able to open the source yet, not that the quotation is wrong.

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