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Origin of the terms embryo, gamete and zygote

Published online by Cambridge University Press:  18 July 2012

Gonzalo Herranz
Gonzalo Herranz*
Affiliation:
Department of Medical Humanities, School of Medicine, University of Navarre, Apartado 177, 31080 Pamplona, Spain.
*
All correspondence to: Gonzalo Herranz. Department of Medical Humanities, School of Medicine, University of Navarre, Apartado 177, 31080 Pamplona, Spain. Tel: +34 948 425 600 ext. 6354. Fax: 34 948 425 630. e-mail: gherranz@unav.es
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The scope of this article is to answer two questions: first, when were the terms embryo (E), gamete (G) and zygote (Z) coined, and second, who were the scientists who put them in circulation? In order to clarify and document the origin of these three words, a vast amount of biomedical literature, particularly from the late 19th and early 20th centuries, has been reviewed; an undertaking made possible by the free access to the massive repository of available scientific literature on the Internet. Furthermore, as a result of this analysis, I would like to raise some ethical aspects of scientific publication, in particular the obligation of scientists to personally assess the quality and source of data they take from the literature and use in their publications.

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Research Article
Information
Zygote , Volume 20 , Issue 4 , November 2012 , pp. 313 - 320
Copyright
Copyright © Cambridge University Press 2012

Origin of the terms embryo, gamete and zygote

The scope of this article is to answer two questions: first, when were the terms embryo (E), gamete (G) and zygote (Z) coinedFootnote 1, and second, who were the scientists who put them in circulation? In order to clarify and document the origin of these three words, a vast amount of biomedical literature, particularly from the late 19th and early 20th centuries, has been reviewed; an undertaking made possible by the free access to the massive repository of available scientific literature on the Internet. Furthermore, as a result of this analysis, I would like to raise some ethical aspects of scientific publication, in particular the obligation of scientists to personally assess the quality and source of data they take from the literature and use in their publications.

The terms E, G and Z originate from the Greek language: E (εμβρυου) is formed from the prefix εν – (in) and the verb βρυειν (to swell, to grow), meaning ‘something that grows inside the body’.Footnote 2 According to the Oxford English Dictionary (OED), G comes from the verb γαμειν (to marry) and its derivatives γαμετ (a wife) and γαμετησ (a husband); and Z derives from ξυγωτοσ (yoked), from the verb ξυγουν (to yoke).Footnote 3 However the history of these words shows interesting differences.

Although E is a very old word, its origin is easy to track. It was used in the distant past by poets and theologians, and was imported into the scientific language at a very early stage. Its history has been relatively uneventful for a long time, but, with the advent of assisted reproduction, it has had a surge of ethical significance and menaced with losing its established profile. By contrast, the search for the origin of the terms G and Z have not been easy. Both were coined together not so long ago by a scientist with the intent of reducing the then dominant lexical complexity, but, regrettably, the memory of the circumstances of place and date of their birth faded away and became lost. To cover the gap, some apocryphal statements on the subject were proposed, that, after persistent repetition in books, journals, and the pages of the web have reached the apparent status of consolidated truth. But, as it will be shown, such attributions are groundless and should be rejected. They constitute one of the strangest fictions in contemporary biological science.

I have arranged this article according to the following plan. First, the data related to the introduction of E in the scientific literature will be discussed. Second, the information gathered from the few dictionaries and encyclopaedias on the coining of the term G will be presented, together with the story of its false attribution to Gregor Mendel. Third, the respective findings related to the term Z, and its erroneous ascription to William Bateson will be presented. Fourth, pertinent data will be shown demonstrating Eduard Strasburger as the creator of the terms G and Z, and 23 November 1877 as the date of birth of both terms. And finally, the weak immediate repercussion that the new terms G and Z received among the scientific community will be alluded to.

I. The past and present of the term E

It is generally agreed that the term E was imported into the scientific literature from the common language. Probably its most ancient use, as early as the 8th century b.c., is found in Homer's OdysseyFootnote 4, when it is applied not to the progeny growing in the womb, but improperly to the already born offspring.

The term then entered the medical language by the hand of the Hippocratic Corpus. In three of its treatises, the word is used with the same meaning as we give it today.Footnote 5 E appears also in many authentic works of GalenFootnote 6, as well as in some spurious books attributed to him. In a medical dictionary it is said that ‘Galen remarks that the Greeks did not call the fetus under 2 months old by the name of embryon, but named it cyema; but others styled it embryon during the whole time of its being in the womb’.Footnote 7

Obviously, with the rediscovery during the late Middle Age of the old Greek medical knowledge, the use of word E was greatly expanded, particularly in the growing literature dealing with human reproduction. The OED points out as the first use of E that of Glanville, De Proprietatibus Rerum, circa 1350, but it is a mistaken attribution.Footnote 8 An unknown disciple of Albertus Magnus authored a widely read book entitled De Secretis Mulierum, its first chapter being entitled De Generatione Embryonis.Footnote 9 It is suggested that this work was composed in the late 13th or the early 14th century. Thenceforth, the use of E became very common.

Some interesting insights on the meaning of E as compared with the word ‘fetus’ for describing different stages of prenatal life are offered by Biggers, who identifies four different ways of using this pair of terms: (as synonymous words, as categorically successive phases of prenatal life, as identifying a subclass of Es called fetuses, and as identifying a subclass of fetuses called Es).Footnote 10 Such usage presupposes a confounding polysemia of the term E. Such a regrettable multiplicity of meanings is inducing a decline of E as a robust biological term: on the one hand, it is being substituted progressively by the more precise words describing its different developmental stages (Z, morula, blastocyst, etc.); on the other hand, after the introduction of the biased neologism pre-embryo, E is losing an important portion of its life span. As anyone has known well, the use and meaning of the term E have been in recent years the focus of intense debates and redefinitions.

II. The supposed origin of the word G

The dating of the G term in dictionaries

The authoritative OEDFootnote 11, which includes among its purposes to record the first time an English word has been used, affirms that the term G made its appearance in 1886. It is attributed to the botanist S.H. Vines in a lengthy article on the reproduction of plants.Footnote 12 Vines used the term G profusely in a bookFootnote 13 published also in 1886, but no claim is made by the author on the coining of G. After 125 years, the attribution to Vines remains unmodified in the OED, despite the fact that a prestigious medical dictionary assigned the introduction of the term G in 1885 to W.R. McNab.Footnote 14

The information given by modern dictionaries available on the Internet on the time of first use of the term G seems to gravitate in the orbit of the OED: with increasing, although only apparent, accuracy they tell us that G was used in the 19th centuryFootnote 15, in the time period ranging from 1885 to 1890Footnote 16, or just in 1886.Footnote 17

The curious attribution to Mendel

In two books specifically devoted to explain the origin of biomedical terms, it is stated that the term G was coined by Mendel .Footnote 18 Skinner says simply the G was introduced by Mendel. According to Haubrich, ‘the biological use of G was advanced by Gregor Johann Mendel (1822–1884), an Austrian monk who gained fame as the naturalist who discovered the fundamental principles of genetics’. Neither Skinner nor Haubrich, however, report when or in which publication Mendel introduced the term G. It is not difficult, however, to ascertain that this is a mistake: simply, the word G does not appear in the incredibly short biological work published by Mendel.Footnote 19

Despite its apocryphal character, this misattribution is disseminated widely in the scientific literature. The phrase ‘the word G was introduced by Austrian biologist Gregor Mendel’ is literally repeated on dozens of websites. Not surprisingly, it has been included in booksFootnote 20, as well as in different versions of Wikipedia.

I have not been able to locate the origin of the false attribution to Mendel of the term G: as happens with many legends, ancient and recent, an anonymous author released it, and many others contributed to its spread. At present, only a few deny it, sometimes for invalid reasons. Such is the case of Bishop, who after declaring that ‘Mendel did not use the term G’, what is true, he adds, apparently by intuition, ‘this term is of the twentieth century’, which is a blunder.Footnote 21

III. The supposed origin of the word Z

The questionable internal consistency of the OED

The OED dates in 1891 the first use of the word Z, assigning it to Hartog.Footnote 22 This is an unfortunate attribution: it cannot be accredited to Hartog as he is not the author of the referenced article. The reference belongs to an anonymous report, included in the ‘News’ section, devoted to summarize the papers read at the Biology Section of the British Association meeting (1891); among them, there is a proposal by Hartog for a new classification of the types of sexual and asexual reproduction. It would be a more fortunate choice to use another sentence appearing above in the same page.Footnote 23

The date (1891) of this attribution seems a little belated and prompts the suspicion that it is in all probability a mistake. Oddly, the OED itself acts as a witness for such a lapse. In its entry G there is a bibliographic reference indicating that 4 years earlier, in 1887, the word Z was present in the scientific languageFootnote 24; in 1891, indeed, it was already frequently used as Hartog himself shows: in a lengthy article published in December of the same year, he uses the term Z 63 times.Footnote 25

In the OED a related term ‘zygosis’ appears as in use 11 years earlier, in 1880. In the corresponding entry, the term is not defined, but its coining is attributed to Pascoe.Footnote 26 The word had a short but interesting life.Footnote 27

The false attribution of Z to Bateson

In parallel to what happened with the false attribution of the word G to Mendel, we find that in virtual or printed texts, the following repeated phrase appears dozens of times and with slight variations ‘William Bateson coined the term Z’. Usually, this false attribution appears included inside a list, otherwise correct, of neologisms created by Bateson, such as geneticsFootnote 28, allelomorph, homozygous and heterozygous.Footnote 29 The dissemination in the literature of the false attribution of Z to Bateson is extensive: it is found everywhere, from books for children30 to well documented on-line encyclopaedias.Footnote 31 This false attribution is not a recent invention: in 1980, the error already was present in some publications.Footnote 32 The mistake is sometimes presented so persuasively that the reader accepts it without further discussion: Henig does just that in her fascinating biography of Mendel.Footnote 33

Indeed, Bateson had used these neologisms (allele, homozygous and heterozygous) in the First Report he wrote, in collaboration with Miss E.R. Saunders, in 1901 on behalf of a Committee for the Royal Society.Footnote 34 It is also true that Bateson used Z and G in 1902, when he voiced his ‘discovery’ of Mendel's workFootnote 35; and also when later he published a more mature presentation of Mendel's findings.Footnote 36 But Bateson never claimed for himself the invention of the word Z. And what is more convincing: he occasionally expressed a certain disdain for such expression, something obviously incoherent with the appreciation authors naturally tend to show for the terms and ideas created by them. In a lecture delivered in 1917 and directed to a lay audience, Bateson stated that the words G and Z are unfortunate expressions. He said ‘They were introduced, according to the haphazard fashion with which scientific men commonly follow in naming their creatures, to denote on the one hand, the single, unpaired germs or germ cells, and on the other the double or paired bodies which result from the union of two germ cells in fertilization.’Footnote 37And immediately he added: ‘The bodies of men and of most of the other animals and plants with which unprofessional people are acquainted are all Z, the product, that is to say, of two G, ‘marrying,’ or germ cells, ‘yoked’ together in a common system, the body.’Footnote 38

But the fact is that the attribution to Bateson has taken deep roots into educational materials intended for information of selected people as developed by institutions of high reputation, such as the Cold Spring Harbor LaboratoryFootnote 39 and the American Philosophical Society.Footnote 40

IV. The birth of Z and G: Eduard Strasburger, Botanische Zeitung, November 23rd, 1877

After an extensive search, a compelling conclusion is clear: both terms, Z and G, were born the same day at the same place. Eduard Strasburger, then Professor of Botany at the University of JenaFootnote 41, introduced them for the first time in the scientific literature, in the issue dated 23 November 1877 of the weekly magazine Botanische Zeitung. The author says in the last column of his article: ‘In conclusion, [. . .] I recommend the name of gametes for these mobile elements[. . .]. The product of the mating of gametes would carry the name of a zygote.’Footnote 42

As the just cited bibliographic reference shows, the work appears to be the result of the cooperation of two co-authors: Anton de Bary and Eduard Strasburger. But, actually, the full and exclusive merit of the creation of G and Z corresponds to Strasburger. A clarification on this point seems necessary.

The cited article appeared in three successive weekly issues of Botanische Zeitung. The journal did not number its pages, but its columns. de Bary is the author of the columns 713 to 728, published in the issue 45 (9 November 1877) and of the columns 729–736 in issue 46 (16 November 1877). Strasburger wrote the columns 737–743 of this same issue 46, and columns 745–758, in next fascicle 47 (dated the 23 November 1877).

The contributions of the two authors are simply juxtaposed: we are not before a joint endeavour, a co-authored piece implying an exchange of data and opinions between the authors, but the product of two independent authors, each of whom wrote his part in the first person singular, put his signature to it and dated it (one in Strassburg, the other in Jena, both in June 1877), and then published both one after the other but under a common title and an apparent co-authorship. It is clear that each author prepared his contribution separately, in ignorance of what the other author did, and, as previously agreed, treated different aspects of the subject matter. There are no cross-references between both articles.

Consequently, each author is the exclusive intellectual owner of the part signed by him. This is recognized by de Bary, at the end of his contribution: he says that by the end of 1876, he and Strasburger agreed to publish, once they had completed some investigations under way, their observations on Acetabularia mediterranea. ‘For publication these conditions were to be followed: although, of course, each knew the main results of his colleague, each would draft his contribution separately, without knowing the manuscript of the other. For my part [de Bary], I would not cite anything that was not already included in my own previous observations, and I would use the terminology then familiar to me. New additions and new findings were to appear in the article by Strasburger that immediately follows.’ Then, de Bary signs and dates his article.Footnote 43

Strasburger, for his part, is aware of the fact he is creating new terminology. He confirms and explains his intentions more clearly in a paper published almost immediately, entitled Über Befruchtung und Zelltheilung [On Fertilization and Cellular Division]. As was then a frequent practice, the paper appeared in two versions: first, in December 1877, as a journal articleFootnote 44; and soon after, in 1878, as a separate book.Footnote 45 The only difference between them is the numbering of the pages. Both, article and book, have a prologue, dated in Jena, in early August 1877, that is, about 2 months after finishing in June the article on Acetabularia sent to Botanische Zeitung, but 3 months before the appearance of the printed article in November 1877. In Über Befruchtung und Zelltheilung, Strasburger offers one reason for the new terminology: ‘The product of conjugation I will name Z, to avoid that the word can be combined with ‘spore’. The Z corresponds to the product of union of egg and sperm and, therefore, ought never to be named spore, as this name must be applied to asexual reproductive cells and therefore restricted exclusively to them. I will call Gs the protoplasmic masses that bind together one another.’ (In a footnote, the author remarks that this matter is addressed in more detail in an article of immediate publication in Botanische Zeitung.) In Über Befruchtung und Zelltheilung, the author used frequently the new two terms.

Obviously, Strasburger does not resort to arguments of authority or indulges himself in explanations of etymological or lexicographical nature. He just recommends the new terms as more appropriate. His purpose is not to create a definitive nomenclature to be used in the future, but to simplify the complex terminology reigning at the moment.

V. The immediate response to the new terms G and Z

Strasburger's message went practically unnoticed. Hardly anyone followed it immediately. De Bary himself disregarded it: he never used the term Z in his publications, and apparently employed G only onceFootnote 46, a fact that could be taken as an indirect confirmation that de Bary had nothing to do with the creation of G and Z.

The reception by scientists of the terms G and Z was a slow business. The echo of both neologisms, especially of Z, in the contemporary biological press was barely audible. In 1878, Strasburger's article was mentioned in two publications: in the section of Notes and Memoranda of the organ of the Society of MicroscopistsFootnote 47, and in The Academy, a weekly high disclosure general (literature, science and arts) at the University of Oxford.Footnote 48

Some scientists welcomed the new words and recognized their usefulness.Footnote 49 Others accepted it only in part or with some reservations.Footnote 50 The dominant attitude was to ignore the new terms. Even Bennett and Murray hold as invalid the reasons given by Strasburger for the changes in terminology, and proposed formally to forget them and return to a more complex system.Footnote 51

G and Z made at last their way to scientific parlance. Julius Sachs used them extensively in his influential book of plants.Footnote 52 Obviously, G and Z had to co-exist with other synonymous terms before becoming the dominant and usual terms, a condition they had already reached in the early years of the 20th century.Footnote 53

Strasburger's attitude toward his neologisms

In the years following 1878, Strasburger used with great restraint, the two words he had coined. For example, in the third edition of one of his most important books, Zellbildung und Zelltheilung Footnote 54 (not to confuse with Über Befruchtung und Zelltheilung), published in 1880, and contrary to what one could expect, the term Z appears in it twice, and G only one. The same happens with an article on the same subject published in 1879 in two successive issues of Botanische Zeitung.Footnote 55 For the rest of his life, Strasburger was remarkably restrained in the use of both words, especially Z, as shown in his studies on reproduction of flowering plantsFootnote 56 and in his collections of histological investigations.Footnote 57 Something similar occurs with the early editions of his famous Treaty of Botany.Footnote 58 In his later years, Strasburger only occasionally reminds his readers that he was the creator of the term G, but strangely practically never proclaims he begot the term Z.Footnote 59

References

Notes

1 For the plural forms (embryos, gametes and zygotes), the abbreviations Es, Gs and Zs will be used.

2 http://www.word-origins.com/definition/embryo.html

3 Simpson, J.A. & Weiner, E.S.C. eds. (1989). The Oxford English Dictionary, 2nd edn.Oxford: Clarendon PressGoogle Scholar.

4 Homer. The Odyssey. For the Greek text: Montgomery, M. (1882). Homer. Odyssey, Book IX. Dublin: Brown & Nolan. The term E appears three times, in the Book IX, verses 245, 309, and 342. Curiously, it is used improperly: it is evident that Homer is not speaking of the unborn in the womb, but of the newborn lamb. See: Crusius, G.C. (1841). Vollständiges Griechisch-Deutsches Wörterbuch über die Gedichte des Homeros und der Homeriden. Hannover. Hahnschen Hofbuchhandlung: p. 156.

5 For example, in the treatises On regimen, On generation, and On the nature of the child.

6 See: Needham, J. (1934). A History of Embryology. Cambridge: University Press: pp. 51–6Google Scholar.

7 Parr, B. (1819). The London Medical Dictionary. Vol. I. Philadelphia: Mitchell, Ames, and White: p. 599Google Scholar.

8 The author of De Proprietatibus was not Bartholomaeus de Glanvilla, but Bartholomaeus Anglicus, who composed this outstanding encyclopaedic book around 1240.

9 (Pseudo) Albertus Magnus. (1655). De Secretis Mulierum. Amstelodami: Iodoam Janssonium; pp. 11–22.

10 Biggers, J.D. (1990). Arbitrary partitions of prenatal life. Hum. Reprod. 5, 1–6CrossRefGoogle ScholarPubMed.

11 The Oxford English Dictionary (see note 3).

12 Vines, S.H. (1886). Reproduction of Plants. Encyclopaedia Britannica, 9th edn, vol. XX. New York: Ch. Scribner's Sons. 425Google Scholar, col. 1.

13 Vines, S.H. (1886). Lectures on the Physiology of Plants. Cambridge: University PressCrossRefGoogle Scholar.

14 Foster, F.P. (1892). An Illustrated Encyclopedic Medical Dictionary, vol. III. New York: Appleton and Co.Google Scholar In the entry G, it is stated: ‘One of two similar or dissimilar reproductive cells, which by their fusion produce a new organism.’ [B, 77, 291 (α, 24); McNab, W.R. Sci. Proc. Roy. Dublin Soc., July, 1885, p. 451].

15 http://www.myetymology.com/english/gamete.html

16 http://dictionary.reference.com/browse/gamete

17 http://www.merriam-webster.com/dictionary/gamete

18 Skinner, H.A. (1976). The Origin of Medical Terms. 2nd edn, reprinted. New York: Hafner Pub. Co.Google Scholar; and Haubrich, W.S. (2003). Medical Meanings: A Glossary of Words Origins. 2nd edn.Philadelphia: American College of PhysiciansGoogle Scholar.

19 In fact, Mendel published only two articles on biological (botanical) subjects: Mendel, G. (1865). Versuche über Pflanzenhybriden. Verhandl Naturf Verein Brünn, 4, 3–46Google Scholar; and Mendel, G. (1869). Ueber einige aus Künstliche Befruchtung gewonnene Hieraciumbastarde. Verhandl Naturf Verein Brünn, 8, 26–31Google Scholar. His other three scientific publications deal with meteorological observations. The full list of Mendel's scientific publications, all of them submissions to the Brünn Naturalists Association, appears in the introductory notes Erich Tschermak wrote for a reprinting of Mendel's biological work: Tschermak, E, ed. (1901). Versuche über Pflanzenhybriden. Zwei Abhandlungen (1865 und 1869) von Gregor Mendel. Leipzig: W. Engelmann, pp. 56–57Google Scholar. Mendel, who certainly contributed basic ideas for understanding the concept of what later would be called G and Z, uses the terms and Pollen-Zelle and Ei-Zelle. The terms ‘pollen cells’ and ‘egg cells’, but not G or Z, appear in the faithful translation into English of Mendel's work, made by his admirer William Bateson (Bateson, W. (1902). Mendel ‘s Principles of Heredity: A Defence. With a translation of Mendel's original papers on hybridisation. Cambridge: University PressGoogle Scholar. This book, significantly amended, was republished 7 years later, (Bateson, W. (1909). Mendel's Principles of Heredity. Cambridge: University PressCrossRefGoogle Scholar).

20 For example, Miller, F.P., Vandome, A.F. & McBrewster, J. eds. (2009). Gamete. Mauritius: Alphascript Publishing.Google Scholar, reads: ‘The name G was introduced by the Austrian biologist Gregor Mendel’. And in Romich, J.A. (2009). An Illustrated Guide to Veterinary Medical Terminology, 3rd edn.New York: Delmar, p. 243Google Scholar, it is stated ‘Johann Gregor Mendel was the first to apply the term G in biology to mean sex cells’.

21 http://heybryan.org/school/Biology/notes/02–27-07,%20Mendelian%20genetics.html

22 Reference in OED reads: ‘1891 Hartog in Nature on September 17. 484 PARAGAMY and ENDOKARYOGAMY: gametal or vegetative nuclei lying in a continuous mass of cytoplasm fuse to form a zygote nucleus.’

23 It is this: ‘KARIOGAMY: the union of cells (gametes), cytoplast to cytoplast and nucleus to nucleus, to form a 1-nucleate cell, the zygote.’ Anonymous. (1891). Biology at the British Association. Nature 44, 481–4CrossRefGoogle Scholar, at 484.

24 This reference says: ‘1887 tr. De Bary's Fungi 495, Gamete, sexual protoplasmic body. . .which in conjugation with another gamete of like or unlike outward form gives rise to a body termed zygote. Same as conjugation-cell.’ In this way, the OED itself moves the birth of the term zygote from 1891 to 1887, and the credit for the word from Hartog to de Bary. But such attribution to de Bary is again a mistake. Only once de Bary used in the original German edition of his book (Vergleichende Morphologie und Biologie der Pilze, Mycetozoen und Bacterien. Leipzig: Engelmann, 1884) the term G (p. 139); but never the word Z. The OED refers logically to the English translation by I.B. Balfour of de Bary's book (Comparative Morphology and Biology of the Fungi, Mycetozoa and Bacteria. Oxford: Clarendon Press, 1887). The quotation in the OED is not de Bary's; it was written by the translator. In effect, Balfour, to help his readers to understand the complex terminology employed by de Bary, added to the book an appendix entitled Explanation of Terms. In the entry for G he says that the conjugation of Gs ‘gives rise to a body termed Z’. Curiously, in the Explanation of Terms there is no entry for Z. In the German original, de Bary used consistently the equivalent term zygospore.

25 Hartog, M.H. (1891). Some Problems of Reproduction: A Comparative Study of Protoplasmic Gametogony and Senescence and Rejuvenescence. Quart. J. Microsc. Sci. 33, 1–79Google Scholar. Hartog offers the following definition of Z: ‘In my terminology I have used the word[. . .]Z to designate the cell produced by their union [of Gs]’. Curiously, the article's recommended reading list includes two books of Eduard Strasburger (‘Befruchtung und Zelltheilung,’ Strasbürger [sic], undated; and ‘Neuere Untersuchungen,’ Strasbürger [sic], 1884).

26 The reference in the OED says: ‘1880 Pascoe. Zool. Chem. [sic] (ed. 2) p. 297’. It corresponds to: Pascoe, F.P. (1880). Zoological Classification: a Book of Reference, 2nd edn.London: Van VoorstGoogle Scholar. The OED's reference to p. 297 directs to the Glossary, although the word zygosis appears in p. 5 of the text, where it is described as ‘one mode of reproduction[. . .]by conjugation’.

27 The concept of zygosis took on a special meaning in the early years of the 20th century. As an anticipation to the debates on sociology and psychology of gender held many decades later, some scientists considered the expressions fertilization and fecundation inappropriate both biologically and sociologically, because those terms were expression of an archaic concept of development, and conveyed the idea, culturally unacceptable, of a passive female G fecundated by an active male G. Therefore, they feel preferable the use of zygosis to denote the fusion of Gs to form the Z, because it described neutrally the phenomenon and was free of social implications. Besides zygosis (suggested by Lankester), other words were proposed then to replace fertilization or fecundation: syngamy and amphimixis recommended respectively by Hartog and Weismann. Each of those neologisms followed different fates. See: Poulton, E.B. (1908). Essays on Evolution 1889–1907. Oxford: Clarendon Press; pp. 60–61Google Scholar.

28 There is general agreement that the noun ‘genetics’ to designate the science of heredity and variation in living organisms was first used by William Bateson in a letter to Adam Sedgwick, dated 18 April 1905. The text of this letter appears in: Bateson, B. ed. (1928). William Bateson, Naturalist. His Essays & Addresses. Cambridge: Cambridge University Press, p. 93Google Scholar.

29 What Bateson did with these last two words was simply to adapt the then already current term Z to prefixes reflecting the new way of understanding Mendelian inheritance. See: Schwartz, J. (2008). In Pursuit of the Gene. From Darwin to DNA. Cambridge, Mass: Harvard University Press, p. 114CrossRefGoogle Scholar.

30 Bardoe, C. (2006). Gregor Mendel: The Friar Who Grew Peas. New York: Abrams BooksGoogle Scholar.

31 In one of them, which allegedly ‘ensures accurate factual information’, we read that ‘In 1905, the English geneticist William Bateson (1861–1926) coined the term ‘genetics’, along with other descriptive terms used in modern genetics, among which include ‘allele (a particular type of gene)’, zygote (fertilized egg), ‘homozygosity’[. . .]. The History of Genetics – Genetics at the Dawn of the Twentieth Century. http://www.libraryindex.com/pages/2216/History-Genetics-GENETICS-AT-DAWN-THETWENTIETH-CENTURY.html

32 In his biography of Gregory Bateson, Lipset notes that the Gregory's father, William, had coined the term ‘genetic’ and introduced the word ‘zygote’, ‘homozygous’ ‘heterozygous’ and ‘allelomorph,’ later shortened to ‘allele ‘(Lipset, D. (1980). Gregory Bateson. The Legacy of a Scientist. Englewood Cliffs, NJ: Prentice-Hall Inc.Google Scholar). Lipset himself, 25 years later, reiterated that William Bateson, who became ‘Mendel's vehement advocate in British biology’, introduced the new terminology it demanded. Genetics, for the new science as a whole, but also Z, homozygote, heterozygote and allelomorph, for the internal relationships between genes.’ Lipset, D. (2005). Author and Hero – Rereading Gregory Bateson: The Legacy of a Scientist. Anthropol. Quart. 78, 899–915CrossRefGoogle Scholar.

33 Henig, R.M. (2000). The Monk in the Garden: Lost and Found Genius of Gregor Mendel, the Father of Genetics. Boston: Houghton Mifflin Co.Google Scholar (The book appeared simultaneously in the United Kingdom with another title: Henig, R.M. (2000). A Monk and Two Peas: The Story of Gregor Mendel and the Discovery of Genetics. London: Weidenfeld & Nicolson.Google Scholar) After referring to Bateson's invention of the term ‘genetics’ says Henig (pp. 229–230): ‘Bateson had already introduced four new terms, which have now been resurrected: ‘zygote’ – from the Greek zugotos, meaning ‘yoked’ – to describe the fertilized egg, the organism formed by the yoking together of two gametes; ‘homozygote’ (from the Greek homos, meaning ‘the same’) and ‘heterozygote’ (from heteros, meaning ‘different’) to describe, respectively, purebreds and hybrids; and ‘allelomorph’ (a compound of allelen, meaning ‘of one another’, plus morphe, meaning ‘form’) to describe the different versions of a particular trait.’ Henig states in a note that her ‘information on terminology is taken from Carlson, Elof Axel (1966). The Gene: A Critical History. Philadelphia: W.B. Saunders.Google Scholar But Carlson did not attributed to Bateson the invention of the term Z. He only transcribed some parts of the First Report to the Royal Society by Bateson & Saunders (see next note 34).

34 Bateson, W. & Saunders, E.R. (1902). The Facts of Heredity in the Light of Mendel's Discovery. Report I, presented Tuesday, December 17, 1901. Reports to the Evolution Committee of the Royal Society. London: Harrison & Sons: pp. 125–160Google Scholar. 126. A verbatim transcript of the report can be found in http://www.esp.org/foundations/genetics/classical/holdings/b/wb-02b.pdf

35 Bateson, W. (1902). Mendel's Principles of Heredity: A Defence. With a Translation of Mendel's Original Papers on Hybridisation. Cambridge: University PressGoogle Scholar. In a footnote on p. 18, Bateson warns: ‘The term ‘G’ is now used as equivalent of ‘germ cells’, and the term ‘Z’ is used here for brevity to refer to the body resulting from fertilization.’

36 Bateson, W. (1909). Mendel's Principles of Heredity. Cambridge: University PressCrossRefGoogle Scholar.

37 Bateson, W. (1928). Gamete and Zygote. A Lay Discourse (The Henry Sidgwick Memorial Lecture, 1917). In: Bateson, B. (1928). William Bateson, Naturalist. His Essays & Addresses Together with a Short Account of His Life. Cambridge: Cambridge University Press, p. 201Google Scholar.

38 In 1909, Bateson gave the term Z a new meaning. He designed as Z both the embryonic and the adult developed, through somatic cell divisions, from the cell resulting from the fertilization process. See: Bateson, W. (1909). Mendel's Principles of Heredity. Cambridge: University Press; p. 16CrossRefGoogle Scholar.

39 Bateson is recognized as the creator of the term Z in the Biography: William Bateson (1861–1926) offered by the Cold Spring Harbor Laboratory's Dolan DNA Learning Center. http://www.dnalc.org/view/16206-Biography-5-William-Bateson-1861–1926-.html.

40 In an introductory note to the collection of writings by William Bateson archived in the Library Association, it is said that ‘[Bateson] left his indelible mark in coining a lot of terminology related to modern Mendelian genetics, from allele to zygote, including the same term ‘genetics’.’ American Philosophical Society. William Bateson Collection 1902–1921. http://www.amphilsoc.org/mole/view?docId=ead/Mss.B.B319-ead.xml; query =; brand = default

41 For a short and informative biography of Strasburger, see: Finke, H.M. Eduard Strasburger – eine biographische Skizze. In Finke, H.M. & Moltmann, U.G. (1994). Hundert Jahre Strasburgers Lehrbuch der Botanik für Hochschulen. Stuttgart: G. Fischer, pp. 29–53Google Scholar. More accessible are, for example, Schwarz-Weig, E. Life and Work of Eduard Strasburger (1844–1912). http://www.deutsche-botanische-gesellschaft.de/html/047StrasburgerCV.html; and, in German Karsten, G. (1912). Nachrufe: Eduard Strasburger. Ber. Dtsch. Bot. Ges. 30, 61–86Google Scholar, The virtual version can be reached at: http://www.archive.org/stream/berichtederdeuts30deut #page/n781/mode/2up

42 deBary, A. Bary, A. & Strasburger, E. (1877). Acetabularia mediterranea. Bot. Zeit. 35, col 713–28Google Scholar, 729–43, and 745–58. The quoted passage appears in column 756. In fact, as shown below, the correct reference ought to be: Strasburger, E. (1877). Acetabularia mediterranea. Bot. Zeit. 35, 745–58Google Scholar, col 756. The publication date is 23 November 1877. The German original reads so: ‘Zum Schluss erlaube ich mir aber, da nun ihr ganzes Verhalten für Acetabularia klar vorliegt, für diese Schwärmer den namen Gameten vorzuschalagen[. . .]. Das Produkt der Paarung der Gameten könnte aber den Namen Zygote führen.’

43 deBary, A. Bary, A. (1877). Acetabularia mediterranea. Bot. Zeit. 35, col 713–28Google Scholar, 729–35, en col 735.

44 Strasburger, E. (1877). Über Befruchtung und Zelltheilung. Jenaische Zeitschr Naturwiss 11, 435–536Google Scholar.

45 Strasburger, E. (1878). Über Befruchtung und Zelltheilung. Jena: Verlag von Hermann Dabis.

46 See note 16, above.

47 In the interest of the terminology introduced, specifically, of the terms of G and Z, the article by de Bary & Strasburger is referred to in: Anonymous. (1878). Notes and Memoranda: Spore Nomenclature. J. Roy. Micros. Soc. 1, 372Google Scholar. But, surprisingly, it is not a direct reference, but only a citation through a secondary source (the Bulletin of the Botanical Society of France).

48 Anonymous. (1878). Science Notes. Botany: Acetabularia mediterranea. The Academy, A Weekly Review of Literature, Science, and Art 13, 79Google Scholar. This note has interesting information: according to it, the contribution of de Bary to the combined article on Acetabularia remained unpublished for 5 years. It mentions that Strasburger calls G to a swarm spore; and Z to the product of fusion of Gs.

49 For example, Fol, H. (1879). Recherches sur la Fécondation et le Commencement de l'Hénogénie chez Divers Animaux. Geneva: Henri Georg, pp. 294–5Google Scholar. Fol, citing the 1877 article in the Jenaische Zeitung, recognized Strasburger as the creator of the terms Z and G.

50 Vines, in his Encyclopaedia Britannica (1886) article, included among its bibliography, a reference to Strasburger's 1878 book Befruchtung und Zelltheilung, and took from it the term G, but he rejected the term Z, and used zygospore instead. Vines, S.H. Reproduction. Encyclopedia Britannica, 9th edn, Vol. XX: p. 425, col 1.

51 Bennet, A.W. & Murray., G. (1880). A reformed system of terminology of reproductive organs of the Cryptogamia. Quart. J. Micr. Soc. 20, 413–20Google Scholar. The authors reject Strasburger's simple new terms and forward instead a cumbersome system. On p. 418, they propose the use of zygosperm to replace Strasburger's Z, and words such as zoozygospheres as substitutes for Gs.

52 Sachs, J. (1882). Vorlesungen über Pflanzen-Physiologie. Leipzig: Vg. Von Wilhelm EngelmannGoogle Scholar. The English translation appeared 5 years later: Sachs, J. (1887). Lectures on the Physiology of Plants. Translated by Ward, H.M.. Oxford: Clarendon PressCrossRefGoogle Scholar.

53 For example, the famous Wilhelm Waldeyer, in a review on the state of knowledge on the sex cells, felt it necessary to clarify the meaning of the Termini technici applied to the reproductive processes in plants and, consequently, to simplify the terminology. He gave preference to G and Z, while acknowledging the usefulness of some terms as macrogamete, microgamete or zygospore. See: Waldeyer, W. Die Geschlechtzellen. In: Hertwig, O., ed. Handbuch der vergleichende und experimentelle Entwickelungslehre der Wirbeltiere. 1. Bd, 1.Teil. Jena: Verlag von Gustav Fischer, 1901–1903 (pp. 388–9)Google Scholar.

54 The book had three editions: Strasburger, E. (1875). Zellbildung und Zelltheilung. Jena: Hermann DabisGoogle Scholar; Strasburger, E. (1876). Zellbildung und Zelltheilung. 2e. Vermehrte und verbesserte Auflage, nebst Untersuchungen über Befruchtung. Jena: Hermann DabisGoogle Scholar; and Strasburger, E. (1880). Zellbildung und Zelltheilung. 3e. Völlig umgearbeitete Auflage. Jena: Verlag von Gustav FischerGoogle Scholar. The 2nd and 3rd editions include as a novelty the study of fertilization.

55 Strasburger, E. (1879). Neue Beobachtungen über Zellbildung und Zelltheilung. Bot. Ztg. 37, col 265–79Google Scholar, 281–8. (Col. 274).

56 Strasburger, E. (1884).Neue Untersuchungen über den Befruchtungsvorgang bei den Phanerogamen als Grundlage für eine Theorie der Zeugung. Jena: Verlag von Gustav FischerCrossRefGoogle Scholar.

57 G and Z are not used in the first of the issues (Strasburger, E. (1988). Histologische Beiträge. 1. Heft. Ueber Kern- und Zelltheilung im Pflanzenreiche, nebst einem Anhang über Befruchtung. Jena: Verlag von Gustav FischerGoogle Scholar), despite the fact, as the title shows, a whole section is devoted to reproduction of plants. In the fourth issue (Strasburger, E. (1892). Histologische Beiträge. 4. Heft. Über das Verhalten des Pollens und die Befruchtungsvorgänge bei den Gymnospermen. – Schwärmsporen, Gameten, pflanzliche Spermatozoiden und das Wesen der Befruchtung. Jena: Gustav Fischer VerlagGoogle Scholar), G is used profusely, and, by contrast, Z only occasionally.

58 Z is quoted once in the first edition, the same as in its English translation: Strasburger, E., Noll, F., Schenck, H. & Schimper, A.F.W. (1894). Lehrbuch der Botanik für Hochschulen. Jena: Verlag von Gustav FischerGoogle Scholar; Strasburger, E., Noll, F., Schenck, H. & Schimper, A.F.W. (1994). A Text-Book of Botany. London: Macmillan & Co.Google Scholar On the history of this book, see: Finke, H.M. & Moltmann, U.G. (1994). Hundert Jahre Strasburgers Lehrbuch der Botanik für Hochschulen. Stuttgart: G. FischerGoogle Scholar.

59 Nearly 30 years after coining the name G, Strasburger notes that ‘the sex cell of single chromosomes receives in plants the name gametophyte, in connection with the general designation of G which I recommended in 1877’[. . .]. Strasburger, E. (1905). Die stofflichen Grundlagen der Vererbung im organischen Reich. Versuch einer gemeinverständlichen Darstellung. Jena: Verlag von Gustav FischerGoogle Scholar; p. 13.