Reprod Dom Anim 47 (Suppl. 4), 2–6 (2012); doi: 10.1111/j.1439-0531.2012.02105.
x
ISSN 0936-6768
An Amazing 10 Years: The Discovery of Egg and Sperm in the 17th Century
M Cobb
Faculty of Life Sciences, University of Manchester, Manchester, UK
Contents
The scientific identification of the key components of sexual
reproduction – eggs and sperm – took place during an
amazing decade of discovery in the 1660s and 1670s. The
names of many of the people involved are now forgotten,
and yet their work, and the difficulties they faced and the
conflicts they endured, resonate strongly to the present day.
Despite this period of innovation, the respective roles of egg
and sperm remained unclear for another 170 years. Why did
this take so long? And what did people think before these
discoveries? By tracing the contours of this major milestone
in human knowledge, we can also gain insight into our
current knowledge, and the boundaries we may be unwit-
tingly trapped by.
Introduction
The 17th century discovery of the role of egg and sperm
in reproduction can be traced to two letters, written
7 years apart, each by a remarkable man who is largely
forgotten today. Those letters heralded an amazing
decade of discovery that eventually shaped the way we
now understand life.
In April 1665, Melchisedec Thévenot (c.1620–92), a
French patron of the sciences, wrote to his friend
Christiaan Huygens (1629–95), a Dutch mathematician
and astronomer: ‘We took the opportunity provided by
the cold of recent months and applied ourselves to
dissections and to investigating the Generation of
animals’ (Thévenot 1665). The ‘we’ referred to two of
Thévenot’s protégés, the Dutchman Jan Swammerdam
(1637–1680) and the Dane Niels Stenson (‘Steno’)
(1638–86). This was the start of a process of discussion,
dissection and experimentation that would soon lead
Swammerdam and Steno to the conclusion that all
animals – including humans – come from eggs.
The second letter was sent 9 years later, in April 1674.
It was written by Henry Oldenburg (c.1615–77), the
German secretary of the Royal Society (Hall 2002) and
was sent to a Delft draper, Leeuwenhoek (1632–1723).
In the letter, Oldenburg asked Leeuwenhoek to use his
microscope to study semen, saliva, chyle, sweat and
other bodily fluids. With this inspiration, in 1677,
Leeuwenhoek would make one of the most stupendous
discoveries in the history of science: the observation of
spermatozoa.
To understand why these two letters were so impor-
tant, we need to unlearn all that we know about
reproduction, beginning with that word. The term
‘reproduction’ was first introduced by Buffon in 1749
(Roger 1997). Up until then, people spoke about
‘generation’, and this was taken to include both how
organisms grow apparently from nothing, and how male
and female contributed to new life (Cole 1930).
Although the simple answer to the question ‘where
do babies come from?’ is fairly obvious – they come
out of the female vagina – arriving at an explanation of
how the baby got there in the first place proved quite
difficult (Cobb 2006a). It seems very likely that early
human populations did not know that intercourse led
to babies. There are number of reasons for thinking
this. Firstly, how could they know? The link between
intercourse and pregnancy is not at all clear or
immediate – people can easily have intercourse without
the woman getting pregnant and the first signs of
pregnancy may not be seen for weeks after the act. This
surprising supposition is supported by the widespread
existence of matrilineal communities in hunter-gatherer
societies, which suggests that men’s role in generation
was uncertain.
It is possible that the domestication of animals
provided the key. In all domesticated animals, mating
takes place only during oestrus (Potts and Short 1999).
Placing the animals together to allow mating would
have been an important step in domestication and in
ensuring the survival of the animals and of the human
group that owned them. According to an un-testable
hypothesis, this could have had two inter-linked conse-
quences: it may have revealed the role of male animals
(and, by extension, of men) in generation, and also
created an economic surplus. And once wealth became
widespread, the question of paternity became funda-
mental for society.
Aristotle, Hippocrates and Harvey
Probably the most influential thinking on generation
came from the Greeks. Aristotle divided all animals into
two kinds – the ‘bloodless’ animals (insects and so on),
which generated spontaneously, and the remainder, in
which mating played a decisive role. For Aristotle, the
woman (or the female animal) provided the ‘matter’ for
the baby, through her menstrual blood, while the male’s
semen gave that ‘matter’ form, like a seal stamping hot
wax. Another analogy, which echoes down to the
present day, was that semen was like a seed (‘semen’
means ‘seed’), which was sowed on fertile ground.
Although the Roman physician Galen adopted Hippo-
crates’ view that there were two ‘semens’ – one male, the
other female – acceptance of this theory was hampered
by the fact that it was not possible to identify the female
semen, and Aristotle’s view predominated. It was also
echoed in the major monotheistic religions of the west
(Judaism, Christianity and Islam), which all stated that
the male semen was the primary component. On the
other side of the planet, Chinese thinking about gener-
ation focused on the ‘generative vitality’ of each sex,
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The Discovery of Egg and Sperm
defined in terms of the energy flows of organ networks,
rather than on particular structures (Furth 1999). On a
world scale, therefore, there was little agreement about
how generation took place or about how each sex
contributed.
In the West (including the Arab world), the ideas of
Aristotle and Hippocrates dominated thinking about
generation for over 1500 years. The first systematic
attempt to explore the problem was made by William
Harvey (1578–1657) in his 1651 book De Generatione
Animalium (On the generation of animals) (Harvey
1651). Harvey was convinced that the ‘egg’ was funda-
mental to generation, although what exactly he meant
by ‘egg’ is unclear. In the 1630s, Harvey dissected red
deer during the rut and tried to find changes in the
females’ ‘testicles’ (as ovaries were then called). But he
could find no changes, nor any sign either of semen or of
an ‘egg’ in the uterus (Short 1978).
Bemused by his failure, Harvey concluded that new
life was produced in the uterus following coitus in the
same way that imagination and appetite are produced in
the brain and that the female’s ‘testicles’ played no role
at all. In the absence of any better evidence, Harvey fell
back on something that looked like Aristotle’s ideas:
eggs were generated like thought, while semen had some
indefinable action at a distance. Even one of the greatest
minds of the age, steeped in the new scientific method
that used experimentation rather than logic, could not
crack the problem of generation.
Women have Eggs
Nevertheless, within 25 years of the publication of
Harvey’s book, thinkers throughout Europe were con-
vinced of the ‘egg theory’ and were certain that all
female animals – including women – produced eggs
(Short 1977). This momentous change came about
because of the inspiration and inquisitiveness of Mel-
chisedec Thévenot, a one-time French ambassador who
had his own small group of scientists and experimenters
(Cobb 2006a). During a visit to Leiden University in the
Dutch Republic, Thévenot had met some brilliant young
medical students, including Swammerdam and Stenson
(‘Steno’), whom he invited to his house outside Paris
and encouraged to start thinking about generation.
Thévenot wanted to show the superiority of the exper-
imental method as against Descartes’ recently published
theoretical approach to generation, which was based
entirely on Aristotle’s ideas.
Although neither Swammerdam nor Steno made any
decisive discoveries in their months with Thévenot, from
mid-1665, they both became focused on trying to
understand generation. Swammerdam returned to the
Dutch Republic and initially focused on the generation
of insects (he was a keen entomologist) and through
careful observation and dissection, he came to the
radical conclusion that ‘all animals come from an egg
laid by a female of the same species’ (Swammerdam
1669). Given that nearly 2000 years of Aristotlean
thought, plus a great deal of casual observation,
suggested that insects appeared from nowhere, Swam-
merdam’s statement – complete with empirical proof –
was remarkable.
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3
Swammerdam’s work appeared the year after an
experimental study by the Italian physician Francesco
Redi (1626–97) which concluded that insects came from
eggs, with the striking exception of gall wasps which he
thought were spontaneously generated (Redi 1668). It
seems probable that Redi was indirectly inspired by
Thévenot, too, as in 1666 Steno had travelled to Florence,
where Redi was the physician to the Tuscan court, and the
two men became close friends. While in Florence, Steno
published what turned out to be his most influential
scientific work, Elementorum Myologiae Specimen (A
model of elements of myology), in which he accurately
described the function of the muscles, using both dissec-
tion and mathematical models (Steno 1668). In an
additional part of this work, Steno incidentally founded
the science of geology, suggesting that the ‘vipers’
tongues’ that were found in rocks and which looked like
sharks’ teeth were in fact the teeth of sharks that had been
stranded on hilltops after the Flood. Most decisively,
Steno included a brief nine-page description of a dissec-
tion of a viviparous dogfish. Primed by his anatomical
dissections in Leiden, and by discussions with Thévenot,
Steno drew a comparison between the anatomy of the
viviparous dogfish and of egg-laying rays: ‘having seen
that the testicles of animals contain eggs, and having
noticed that their uterus opened into the abdomen like an
oviduct, I have no doubt that the testicles of women are
analogous to the ovary’ (Steno 1668).
The Race to Find Proof
This simple but revolutionary statement – which had no
proof to back it up – soon caused consternation back in
Leiden, where Swammerdam and Professor Van Horne
(1668) had already set out about trying to identify eggs
in the ovaries of dead women. Van Horne, like his
former pupil, Steno, had come to the conclusion that the
vesicles in the woman’s ‘testicles’ were, or contained,
eggs. Together with Swammerdam, van Horne even
showed that the vesicles in the ovaries of a cow turned
white when boiled, just like a chicken’s egg. But it now
appeared that their friend Steno had published the
discovery first. Their only hope was to provide what
Steno’s brief account lacked: proof. A few months later,
van Horne published a brief summary of his discoveries
(Van Horne 1668), but the major work that he promised
himself and Swammerdam never appeared. In January
1670, van Horne died of a plague that swept through
Leiden, killing five of the University’s 15 professors
(Luyendijk-Elshout 1965).
After van Horne’s death, Swammerdam continued
their work, but soon became aware that someone else
was studying the same problem: his one-time student
friend, de Graaf (1641–73). In May 1671, following a
brief meeting at which it became apparent that they were
both approaching completion, de Graaf hastily pub-
lished a brief outline of his work, in which he summa-
rized his view of how ‘eggs’ in the ‘testicle’ became
‘fertile’ through the action of the ‘seminal vapour’ rising
up from the uterus via the Fallopian tubes (De Graaf
1671). In December 1671, Swammerdam riposted by
publishing a single sheet drawing of his dissection of the
human ovary and the uterus, which he sent to the Royal
4 M Cobb
Society in London (Swammerdam n.d.). Although neither
man had yet provided any clear evidence that human
ovaries contained eggs, they were now on a collision
course.
De Graaf apparently won the race when, in March
1672, he published his brilliant De Mulierum Organis
Generationi Inservientibus Tractatus Novus (New treatise
concerning the generative organs of women) (De Graaf
1672; Jocelyn and Setchell 1972). Although the book
contained dissections of humans, rabbits, hares, dogs,
pigs, sheep and cows, its decisive part was a section on
rabbit mating and pregnancy. Here, de Graaf referred to
the follicles or their contents as eggs, and gracefully gave
his former teacher, van Horne, the credit for this
discovery, pointing out that Steno had merely said that
ovaries contained eggs without identifying them. Above
all, de Graaf used careful experimental dissection to
show that in rabbits the follicles reddened and ruptured
following mating and that 3 days after copulation, small
spherical structures could be found in the Fallopian
tubes. De Graaf emphasized that the number of these
spheres was generally identical to the number of ruptured
follicles and that it never exceeded them. Because mating
induces follicular rupture in the rabbit, de Graaf
mistakenly suggested that the same thing must happen
in women, which would imply that virgins would show
no ruptured follicles. Finally, like Harvey, de Graaf
looked for signs of semen in the uterus and Fallopian
tubes, but could find none. He concluded that only a
‘seminal vapour’ reached the eggs and fertilized them.
Through his experimental work, de Graaf showed the
power of the scientific method, and also demonstrated
that mammals – including women – have eggs.
In response, in May 1672, Swammerdam published
his own account of human generation, Miraculum
Naturae, sive Uteri Muliebris Fabrica (The miracle of
nature, or the structure of the female uterus) (Swam-
merdam 1672a,b). This contained no experimental
studies, merely a great deal of dissection, some of which
was highly pertinent, as it showed that virgin women
also have ruptured follicles. Swammerdam also explored
the very real problem of how the fluid in the follicle –
which he felt was the egg – could move from the ovary
across into the Fallopian tube. But the central theme
was Swammerdam’s claim that van Horne had been the
first person to accurately describe the female reproduc-
tive organs and that he, Swammerdam, had first
suggested that the egg was in the follicle. The book
closed with a polemical appendix, in which Swammer-
dam sneered at the quality of de Graaf’s drawings and
even at the fact he had studied ‘rabbits and brute
animals’ rather than humans, thereby inadvertently
showing that he had entirely missed the point of de
Graaf’s work. Swammerdam dedicated his book to the
Royal Society, and sent it off to London, followed by a
beautifully preserved human uterus, along with twelve
other items of genital anatomy, both male and female.
Above all, Swammerdam asked the Royal Society to
adjudicate on who had the priority in stating that
women have eggs.
Less than a year later, as the Dutch Republic was
wracked by a murderous invasion by the French in the
opening battle of a war that lasted 6 years, de Graaf
produced a blistering response to Swammerdam, Par-
tium Genitalium Defensio (Defence of the genital parts)
(De Graaf 1673a,b). This savage polemic – written in the
immediate aftermath of the death of de Graaf’s month-
old baby son, Frederick – reproduced letters between de
Graaf and Swammerdam, quoted conversations and
made accusations that far exceeded anything seen in a
modern row on the internet. De Graaf flatly denied that
he knew that Steno had ever published about eggs (the
two men had in fact corresponded on the question
several times), and accused Swammerdam of plagiarism
and of piling ‘lie upon lie’. De Graaf then sent what he
admitted was a ‘not very polite’ book to the Royal
Society and, like Swammerdam, asked them to judge
who had priority.
In response to the pleas from the excitable Dutchmen,
the Royal Society set up a three-man committee to deal
with the fractious dispute. When the committee eventu-
ally reported, in October 1673, they decided that neither
de Graaf, nor Swammerdam, nor van Horne had been
first to discover that women have eggs. That honour,
they stated, went to Steno. This verdict turned out to be
completely pointless. A week before the committee
completed its work, de Graaf died, aged only 32.
Swammerdam wrote an immediate reply, of which no
trace remains, and then never referred to the matter
again. Steno, meanwhile, was on the verge of abandon-
ing science to become a Catholic bishop, and there is no
evidence he ever heard of his ‘victory’. Even the Royal
Society seems to have lost heart, for the report was not
published for more than 80 years (Birch 1756–7).
Ironically, history has adopted a very different view:
the follicles are now known as Graafian follicles and the
part played by Steno, Swammerdam and van Horne is
forgotten to all but a handful of historians.
Whatever the ins and outs of the priority dispute, the
key issue was that by the mid-1670s, the ‘egg theory’
came to dominate. In 1679, the French scientific
publication Journal des Sçavans wrote: ‘The view that
man, as well as all other animals, are formed from eggs
is something that is now so widespread that there are
hardly any new philosophers who do not now accept it’
(Anonymous 1679). However, even as the ink was
drying on the page, an amazing new discovery was
challenging that view, and would once again throw the
scientific community into turmoil.
Enter the Sperm
When Reinier de Graaf sent his polemical book to the
Royal Society in April 1673, he included a report which
a friend of his, ‘a certain very ingenious person named
Leeuwenhoek, has achieved by means of microscopes’
(De Graaf 1673a,b). Leeuwenhoek’s brief letter included
descriptions of a bee sting, a louse and a moss. This was
the beginning of a long relationship between Leeuwen-
hoek and the Royal Society, which spanned the next
50 years and consisted of nearly 200 letters from the
Dutchman (Dobell 1932).
Unlike de Graaf, Steno and Swammerdam, Leeuwen-
hoek (he later adopted the prefix ‘van’ as an affectation)
was not academically trained. He was a draper, who had
begun making his own single-lens microscopes for
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The Discovery of Egg and Sperm
reasons that remain obscure (Ford 1985). Although the
microscope had been invented at the beginning of the
17th century, they were generally little better than
magnifying glasses (Ruestow 1995; Wilson 1995). The
power of this instrument was brought to public atten-
tion in 1665 when Robert Hooke (1635–1703) published
his magnificent book Micrographia, complete with
stunning illustrations (the full text of this amazing work
can be found on the internet). But Hooke used a
compound microscope, which was very difficult to focus,
as Samuel Pepys (a member of the Royal Society)
discovered when he bought one of these new-fangled
devices having been impressed by Hooke’s book (‘the
most ingenious book that I ever read in my life’). To his
disappointment, Pepys found it very difficult to see
anything like the images printed by Hooke, and he felt
that the princely sum of £5 10s he had spent on the
microscope was wasted (Tomalin 2002).
The single-lens microscope was far easier to construct
– it simply involved polishing a tiny glass ball approx-
imately 1 mm across – and it was widely adopted in the
Dutch Republic, with some of its main practitioners
including Swammerdam, who saw red blood cells and
drew the first cell divisions in a fertilized frog egg, and
the great philosopher Benedict Spinoza (Ruestow 1995).
Leeuwenhoek’s long career as a pioneer microscopist
was encouraged by Henry Oldenburg, who published
Leeuwenhoek’s first letter in the Philosophical Transac-
tions and then, in 1674, wrote that letter to Leeuwen-
hoek, inviting him to turn his attention to the
composition of various bodily fluids, including semen
(Cole 1930). Over the next few years, Leeuwenhoek sent
a number of letters to London, amazing his readers with
descriptions of tiny ‘animalcules’ in water (these were
protists and large bacteria) (Leewenhoecks (sic) (1677).
Then, in autumn 1677, a young medical student from
Leiden, Johann Ham, brought Leeuwenhoek some pus
mixed with semen to examine. Ham claimed that this
had been produced by a ‘friend’ who had ‘lain with an
unclean woman’. When Ham had looked at the sample
under his microscope, he had noticed many ‘animal-
cules’ in it; alarmed or enthused (it is not clear which),
Ham brought the sample to Leeuwenhoek, who con-
firmed the observation and did the obvious thing: he
looked at his own semen. To his amazement, Leeuwen-
hoek saw there were millions of tiny animalcules
thrashing about in the sample. These things were
eventually given the name by which they are still known
today, and which completely misclassifies them: ‘sper-
matozoa’ – semen animals.
Three months later, Leeuwenhoek dispatched a care-
fully worded letter to London, written in Latin, insisting
that should the society consider it ‘either disgusting, or
likely to seem offensive to the learned, I earnestly beg
that [it] be regarded private and either published or
suppressed as your Lordship’s judgement dictates’
(Leeuwenhoek 1678). In a process that will be familiar
to modern readers, the Royal Society then sat on
Leeuwenhoek’s earth-shattering discovery. Oldenburg
had died a few months earlier, and there was a hiatus in
the offices of the Philosophical Transactions. The new
editor, Nehemiah Grew, had his doubts about Leeu-
wenhoek’s findings and eventually wrote back in Janu-
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5
ary 1678 asking him to look at the semen of dogs, horses
and other animals. Two further letters from Delft
followed with the requested observations, but they
merely added to Leeuwenhoek’s previous findings. The
lack of urgency suggests that no one at the Royal
Society understood why Leeuwenhoek’s discovery was
so important (they do not appear to have considered it
‘disgusting’). Leeuwenhoek’s letter was finally published
in January 1679, nearly 18 months after his initial
discovery (Leeuwenhoek 1678).
Leeuwenhoek’s letter makes clear that he thought that
the spermatozoa were merely another example of the
‘animalcules’ he could see everywhere he pointed his
microscope. He was much more interested in a tangled
mass of tiny vessels which he saw in ‘the denser substance
of the semen’. This structure led Leeuwenhoek to claim
without the slightest evidence that ‘it is exclusively the
male semen that forms the foetus and that all the woman
may contribute only serves to receive the semen and feed
it’. In other words, Aristotle had been right all along. The
exact nature of the mass of vessels remains obscure –
Leeuwenhoek later admitted that it was ‘merely acciden-
tal’ (Leeuwenhoek 1683). Whatever the case, Leeuwen-
hoek did not immediately grasp the significance of his
discovery of animalcules in semen.
Other people were far more astute than either Leeu-
wenhoek or the Royal Society, and realized that the
animalcules themselves were highly significant. Prior to
publication, news got out about what Leeuwenhoek had
seen. A few months after Leeuwenhoek’s original obser-
vation, in January 1678, Swammerdam wrote a letter to
Thévenot stating that he had observed ‘innumerable small
worms’ in mouse and dog semen (Lindeboom 1975). Even
more importantly, in the summer of 1678 – over 6 months
before the Royal Society published Leeuwenhoek’s letter
– Huygens printed a brief account of the study in the
Journal des Sçavans, which concluded: ‘This latest
discovery, which has been made in Holland for the first
time, seems to me to be extremely important and will
provide material for those who seriously study the
generation of animals’ (Huygens 1678).
150 Years of Confusion
A naive modern reader could reasonably assume that
this was the end of the matter and that everyone soon
realized that egg and sperm were complementary, each
containing half of what was necessary to produce new
life. Not at all. Firstly, there were technical issues: no
one had yet seen a human egg, and would not do so until
1827 (Von Baer 1956). But above all, it was not clear
what the discoveries meant. For nearly 150 years,
thinking about generation was dominated by either
‘ovist’ or ‘spermist’ views (Pinto-Correia 1997; Roger
1997). Each approach considered that only one of the
two parental components provided the stuff of which
new life was made, with the other component was either
food (as the spermists saw the egg), or an immaterial
force that merely ‘awoke’ the egg (as the ovists saw the
spermatozoa).
There were many reasons underlying this apparent
scientific dead end. For example, in chickens the two
elements did not seem to be equivalent at all: there was a
6 M Cobb

single, enormous egg which was apparently passive, while
the ‘spermatic animals’ were microscopic, incredibly
active and present in mind-boggling numbers. Ultimately,
however, the reason why late 17th-century thinkers did
not realize what to us seems blindingly obvious – that both
egg and sperm make equal contributions to the future
offspring – was that there was no compelling evidence to
make them appreciate this.
It was not until the 19th century that the requisite
combination of evidence and theory came together. To
understand the complementary nature of egg and sperm,
scientists needed to have a theory that could explain that
complementarity. This came in two forms in the early
decades of the 19th century (Cobb 2006b). The devel-
opment of ‘cell theory’ by Schleiden and Schwann gave
an explanation for why egg and sperm were equivalent,
despite their manifold differences – they were both
reproductive cells. The other factor was that realization
that heredity had a biological content and that some-
thing was inherited, which was contained in egg and
sperm, respectively. This development came about
through the conjunction of three areas: the work of
agro-industrialists such as Robert Bakewell, who carried
out massive selective breeding programmes on domes-
ticated animals; the studies by thinkers such as Mau-
pertuis and Réaumur, who explored large and complex
family trees in the light of particular characters; and by
French physicians who studied the inheritance of
diseases (Cobb 2006b). This was finally given form in
a monastery in Brno, where Gregor Mendel was just one
of many people thinking about the nature of heredity.
Oddly enough, by the time that the fusion of egg and
sperm was observed for the first time, by Hertwig and
Fol in the late 1870s, it was almost an anti-climax.
People thought it was obvious. Which in a way, it was,
but getting to such a point had been anything but
straightforward, and would not have occurred without
that exciting spasm of discovery in the 17th century.
Conflicts of interest
None of the authors have any conflicts of interest to declare.

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Author’s address (for correspondence):
M Cobb, Faculty of Life Sciences, University
of Manchester, Oxford Road, Manchester
M13 9PT, UK. E-mail: cobb@manchester.
ac.uk

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