JOURNAL OF THE ROYAL SOCIETY OF MEDICINE Volume 88 September 1995

Anatomy of a physiological discovery: William Harvey and

the circulation of the blood
Emerson Thomas McMullen PhD

J R Soc Med 1995;88:491-498

Keywords: William Harvey; circulation of blood; valves in the veins

INTRODUCTION Dropping the above phrase with 'recently' in it would not

Historians have dissected William Harvey's conception that have altered at all the point Harvey was making about
the blood circulates in terms of what, how, and when. respiration. Therefore his reference to a 1615 book as
Examples of what they thought initially led to Harvey's 'recent' tends to support the literal meaning of 'nine years
mental discovery include the sameness of the blood1 and and more'. The implication of both quotations is that Harvey
Harvey's intelligent experimental method2. A currently had conceived of the blood's circulation earlier than
popular view centres around the blood's quantity3. As for conventionally thought, and that he was writing about it
how Harvey did it, opinions vary widely. One is that he used before 1619.
an approach based on function and the hypothetico-deductive The last clue in De motu cordis on the time frame of the
method4. When Harvey made his discovery also has been discovery is in Chapter 13. This chapter contains Harvey's
debated. Recent estimates fall between 1619 and 16285. discourse on the valves in the veins and the only pictures in
This article presents evidence that the conception process the book. Here Harvey referred to Fabricius' work on the
began before 1617 and that purpose was more important venous valves, and most interestingly, described his old
than function. Function describes how something works teacher. Harvey portrayed Fabricius as 'a man advanced in
while purpose explains why it is there in the first place. The years'. Harvey did not say 'who recently passed away', or
purpose of a clock, for example, is to tell time, while something similar. Obviously Fabricius was living at the time
function explains the workings of its gears and springs. Harvey wrote this and did not pass off the scene until 21
May 1619. Therefore, the middle of 1619 has to be an
WHEN WAS THE DISCOVERY? absolute upper limit for when Harvey wrote this key
There are several places in Exercitatio anatomica de motu cordis chapter. This early date creates difficulties for some ideas on
et sanguinis animalibus (hereafter called De motu cordis), which
Harvey and his discovery: one such idea is that Harvey wrote
De motu cordis in two stages8.
contain clues to the time frame of Harvey's mental
discovery. The first is well known. In the Dedication to The evidence from Harvey's lecture notebook is that he
Dr Argent, President of the Royal College of Physicians,
had not made the discovery in 1616. This date, then,
Harvey wrote that he had confirmed his new views for 'nine
becomes an absolute lower limit. Probably, Harvey had the
years and more' earlier6. Since De motu cordis was first
complete hypothesis before he started writing9. Thus, a
published in 1628, then 'more than nine years' had to be at possible scenario is that Harvey made the complete
least in 1618, or even earlier. Also, later in the Dedication, breakthrough in 1617 and then immediately wrote the
he stated that his 'little book' was 'complete some years bulk of De motu cordis. (The Dedication was either updated
or else written much later.) If so, then Harvey's rejection of
ago'. This early time frame for the discovery could explain
hints that some of the medical community on the continent the old physiology was even earlier than 1617. Support for
this early rejection comes from a study of the valves in the
heard rumours of Harvey's discovery by 16227.
After the Dedication of De motu cordis is the Introductory veins.
Discourse. The clue to the time frame involves Harvey's old
anatomy professor at Padua. Harvey identified Hieronymus THE VENOUS VALVES
Fabricius' De respiratione as 'the most recently published The early discoverers of the valves in the veins understood
book on the subject'. De respiratione was published in 1615. their function correctly. Amatus Lusitanus (1511-1568)
It is hard to know precisely what Harvey meant by wrote that 'ostiola' (little doors) prevent the blood from
'recently', but he did not have to use the word at all. returning, and operate like those in the heart. Andreas
Vesalius (1514-1564) reported that Giovanbattista Canano
(1515-1579) told him that membranes in the veins, similar
Historian of Science, Technology and Medicine, Department of History, Georgia to those in the heart, prevented the reflux of blood.
Southem University, Statesboro, Georgia, 30460-8054, USA However, Vesalius himself thought these membranes 491
 JOURNAL OF THE ROYAL SOCIETY OF MEDICINE Volume 88 September 1 995

strengthened the veins10, and because of Vesalius' somewhat

negative influence, interest in the venous valves waned. This
situation changed with Fabricius, Harvey's most influential
teacher1l. In 1574, Fabricius claimed to have discovered the
'valves' in the veins. He called them 'ostiola', meaning
'little doors'. (Hereafter, the term 'ostiola' will be used
instead of 'little doors'.) In 1603, Fabricius published the
definitive work on these venous membranes, De venarum
ostiolis12.
A top anatomist, Fabricius might have discovered the
blood's circulation. According to Galen of Pergamon
(second century AD), blood ebbs and flows in the arteries,
distributing the vital spirit. Blood moves similarly in the
veins, nourishing the body, but with a more general outward
flow from the heart as the blood, or part of it, is consumed. Figure 1 From Salomon Alberti, Tres Orationes, 1585, showing the
outside (left picture) and inside (right picture) of part of a leg vein,
The discovery of ostiola unbalanced the physiological with the two cusps of a bicuspid valve in the vein. These are the
symmetry of Galen's explanation. Further, if ostiola acted first drawings of a valve
like check-valves and allowed blood to flow only toward the
heart, then the body's extremities would starve, in which nature. He also reported the early accounts of the venous
case Galen's physiology would be obviously incorrect. valves, including the findings of Lusitanus and Canano.
Fabricius thought that ostiola function not as one-way Alberti had used several terms including 'valvulis' for the
valves, but only as hindrances to the blood's outward flow. venous valves, but Fabricius, consistent with his own
Based on this function, he argued that the purpose of ostiola nomenclature, reported in De venarum ostiolis that Alberti
was to slow the blood's flow, preventing it from collecting had written 'most learnedly on the "ostiola" of veins'. In
too rapidly in the body's extremities. If the blood flowed too his anatomical lecture notes, Harvey referred to Fabricius by
quickly to the hands and feet, other body parts would be name six times, and to Alberti three times. Thus, early on
undernourished. Fabricius drew an analogy between ostiola Harvey was faced with a clear choice between ostiola, as
and floodgates which hinder water flow in the sluices at a described by Fabricius, or the one-way valves in the veins, as
mill. Thus, it would appear that he explained the purpose of described by Alberti, Lusitanus, and Canano. We will see
ostiola in terms of accepted physiology. that Harvey made this choice very early.
In reality, Fabricius modified Galen's paradigm. To keep
the blood from falling down into the lower parts of limbs,
Galen had hypothesized the existence of an 'attractive' THE KEY TO THE DISCOVERY
power. Fabricius replaced that notion with his more Robert Boyle (1627-1691) has been called the 'Father of
mechanical explanation. In this limited way, he had Chemistry' 13, and is well-known for the physical law which
developed a new physiology. Perhaps this is one reason he bears his name. What is not so well known is that he was
ended De venarum ostiolis by saying: intensely interested in medicine, and was awarded a medical
degree in 166514. Boyle authored medical books ranging
such is the wisdom and ingenuity of Nature which by my own efforts from his important Memoirsfor the Natural History of Humane
I have discovered in this new field. Blood, Especially the Spirit of that Liquor (London: 1684), to
several editions of remedies for laymen, Medicinal
Fabricius criticized Vesalius for not going beyond careful Experiments: or a Collection of Choice and Safe Remedies, For
anatomy to physiology. This is why Harvey was such a The most part Simple and easily prepared: Very useful in Families,
student of Fabricius. Vesalius had broken new ground in andfttedfor the Service of Country people (London: 3rd ed.,
anatomy. Fabricius realized that the next frontier would be 1696). Scattered through Boyle's works are so many
in physiology. Fabricius and Harvey did go beyond careful remembrances relating to a conversation he had with Harvey
anatomy to physiology, but Harvey's physiology was more that some scholars think Boyle had several meetings with
careful, and therefore more productive, than that of Harvey, even though Boyle claimed only one15.
Fabricius. Boyle had interviewed Harvey about the discovery of the
Another person writing on the venous valves and the first blood's circulation some 30 years after the event. Boyle
to picture them was Salomon Alberti, a medical professor at delayed publishing the information from this meeting. These
the University at Wittenberg. In Tres Orationes, 1585, he circumstances raise concerns about whether Harvey recalled
492 illustrated the valves (Figure 1) and noted their one-way the true details of the discovery, whether Boyle heard rightly
 i -~ ~ ~ ~ .,:
i i^
JOURNAL OF THE ROYAL SOCIETY OF MEDICINE

. _
Volume 88

of Fabriciust7. Harvey meant it when he wrote in

September 1995

way, like the heart's. Harvey knew Fabricius' work well.

The only pictures in De motu cordis are nearly identical to one

Exercitationes de generatione animalium (1651) that from

among ancients he followed Aristotle, but from among
contemporaries, he followed Fabricius.
The second of Boyle's accounts of Harvey's discovery is
found in A Disquisition About the Final Causes of Natural Things
(1688). Boyle dictated Disquisition in the 1670s to clarify and
extend his earlier comments on purpose. However, Boyle
stated that he never consulted his previous writings.
Therefore, when he came to his Harveian example, he did
not check what he had dictated earlier. Instead, he recalled
the conversation with Harvey from memory. This
recollection differs from what he dictated before:

And I remember that when I asked our famous Harvey, in the

discourse I had with him, (which was but a while before he died),
only
what were the things that induced him to think of a circulation of the
Blood? He answered me, that when he took notice that the valves in
the veins of so many several parts of the body, were so placed that they

Figure 2 Table Ill from Fabriclus' De venarum ostiolis, 1603. Part

of the heart and related vessels. In the upper left (at the origin of
the bifurcated aorta) are the three semilunar cusps of the aortic
valve, labelled F.F.F. In the text, Fabricius points out their close
similarity to the cusps of the venous values such as those depicted
in Figure 3

what Harvey said, and whether Boyle correctly remembered

the interview. All this is compounded by the fact that
Boyle's two accounts of Harvey's discovery differ from each
other. One account is in Boyle's Some Considerations touching
the Usefulness of Experimental Natural Philosophy'6 dictated
around 1650 (7 years before Harvey's death) but not
published until 1663. In this work Boyle reported that 'the.........
structure of the valves of the heart and veins' first influenced
Harvey. Boyle did not say what it was about this structure
that led Harvey to conceive of the blood's circulation.
Somehow, the structure of the venous valves was linked to
those in the heart. ..

The link involving Harvey's discovery was that heart k .

valves allow one-way flow only, and their structure,
particularly the aortic and pulmonary valves, is similar to
that of venous valves. Harvey may have learned of this
simiflarity from Fabricius. In De venarum ostiolis, Fabricius
pointed out how the aortic valve (Figure 2) is like the venous
valves (Figure 3). The only obvious difference besides size is
cusps, while
that the aortic valve has three otiavehsthe
that th uss the venous valves
hieteeosavs Figure 3 Table viii from Fabricius' De venarum ostiolis. The
in a bifurcated leg vein. Fabricius noted the likeness of their valves
have one or two. Basically, these cusps appear the same. and single
double cusps to the triple cusps of the aortic valve as pictured
This comparison suggests that the venous valves are one- in Figure 2 493
 JOURNAL OF THE ROYAL SOCIETY OF MEDICINE Volume 88 September 1995

gave free passage to the blood towards the heart, but opposed the defective valves might not be competent, but so many valves
passage of the venal blood the contrary way: he was invited to result in one-way flow, creating an anomalous situation.
imagine, that so provident a cause as nature had not plac'd so many In this same section of De motu cordis, Harvey also
valves without design: and no design seemed more probable, than that,
since the blood could not well, because of the interposing valves, be rejected Fabricius' explanation because of the orientation of
sent by the veins to the limbs, it should be sent through the arteries, the valves in the jugular veins. He wrote that the discoverer
and return through the veins, whose valves did not oppose its course of the valves 'did not rightly understand' their purpose:
that way [emphasis added]"8.
... for their office is by no means explained when we are told that
In this account, Harvey initially made his discovery by it is to hinder the blood, by its weight, from all flowing into
observing the orientation of the venous valves and asking, in inferior parts; for the edges of the valves in the jugular veins hang
downwards and are so contrived that they prevent the blood from
effect, why so many? rising upwards . . .
Since not all veins have valves, one could argue that the
valves are not that abundant. Harvey did not share this Harvey understood Fabricius' explanation for the valves'
perception: he was struck by their quantity. He twice purpose, but saw no support for it. In this chapter he used
mentioned the abundance of these valves in the above the Latin term 'valvulas' (or a form of it) and not Fabricius'
passage. This led to the implied question of why there are no 'ostiolis' to identify the cusps in the veins. This difference in
valves in the arteries? In contrast to the other Boyleian nomenclature accurately reflects the different role these
account, Harvey initially did not ask about the venous valves cusps have in Fabricius' physiology compared to their role in
or their structure, but about their orientation and relative Harvey's. When and why did Harvey realize Fabricius was
abundance. Some historians have noted the role of the wrong?
venous valves19 or their orientation20 in Harvey's discovery,
but none have pointed out the significance Harvey placed on
their abundance. THE EARLY EVIDENCE
Harvey's stress on both the valves' orientation and their
abundance undercuts the idea that later emphasis on the As mentioned earlier, Salomon Alberti described the one-
structure of the venous valves influenced Boyle's or Harvey's way nature of the valves in the veins and used the Latin term
memory. Harvey's observation of the orientation and 'valvulis' for them. Immediately following the section on the
abundance of venous valves are unique remembrances of venous valves, he wrote about the ileocaecal valve located
the discovery. There was no earlier emphasis on the valves' between the small and large intestines. Alberti used the
orientation and abundance which could have affected Boyle's terms 'valva' and 'valvulas' for this valve. Harvey read this
or Harvey's recollection. This fact alone should make section and in his lecture notebook, Prelectiones Anatomiae
scholars such as J J Bylebyl cautious about discounting Universalis (1616), he compared the function of the ileocaecal
Boyle's report21. Boyle was capable of making mistakes, but valve to that of the valves in the veins:
it was not like him to invent an account out of thin air. The
best explanation is that Boyle did not contradict himself, but WH Those who say as Sal[omon] Alb[ertil that there is within [the
remembered part of the conversation in one treatise and part bowel] a membrane which closes the passage as in the veins22.
in another. This interpretation is supported by references to
the number and orientation of venous valves in Harvey's The context of this passage is that the ileocaecal valve stops
works. reverse flow. (Harvey incorrectly used 'membrane' to
For instance, Harvey discussed the number (and describe the function of this sphincter valve. Since Alberti
competence) of venous valves in Chapter 13 of De motu also had used 'membrane' to describe the venous valves, it is
cordis: obvious that Harvey envisioned a functional connection
between the two types of one-way valves.) Of interest to us
And although in some places the valves, by not acting with such is that Harvey treated the one-way nature of the venous
perfect accuracy, or where there is but a single valye, do not seem valves as a given. He compared the ileocaecal valve to the
totally to prevent the passage of the blood from the centre, still the venous valves and not vice versa23. Further, he was not just
greater number of them plainly do so; and then, where things appear
contrived more negligently, this is compensated either by the referring to what someone else was saying: he denoted his
more frequent occurrence or more perfect action of the succeeding acceptance of this check-valve operation in his customary
valves fashion by putting his initials next to the passage. Therefore,
prior to 1616, Harvey had accepted that the membranous
Where a valve might be 'contrived more negligently', or, valves in the veins basically stop reverse flow. This implies
where there is but a single valve, such situations are that the old explanations of the blood's motion had to be
494 compensated for by the abundance of valves. A few single or incorrect.
 JOURNAL OF THE ROYAL SOCIETY OF MEDICINE Volume 88 September 1995

In addition, Harvey mentioned the abundance and in the arteries (save at their roots) ....' This difference
orientation of the venous valves later in his Prelectiones. from the venous valves is only implied in the Boyle
Writing before he hypothesized the blood's circulation, reports)
Harvey remarked that: 5 All venous valves have the same orientation, ('opposed
to the heart')
WH Wherefore there are many valves in the veins opposed to the
heart; the arteries have none except at the exit from the heart. The broader context of the above passage from
Prelectiones involves the heart's pulse, not the blood's
This important early observation has a lot packed in it. circulation. However, what is important here again is
Analysing the above passage, we find that part of it Harvey's nomenclature. He wrote 'valvulas', not Fabricius'
encapsulates Boyle's reports about the discovery, and part 'ostiola' for the valves in the veins. Later, Harvey would
relates to what Harvey later wrote about the venous valves freely copy Fabricius' picture because it was accurate
in De motu cordis: (compare Figures 4 and 5). In Preleciones, Harvey is not
using Fabricius' terminology because, even at this early date,
1 There is more emphasis on the valves' abundance than he already doubts Fabricius' explanation.
on their structure
2 There is a link to the heart, (the aortic valve 'at the exit
ANATOMY OF A REJECTION
from the heart'. This similarity of the venous and
cardiac valves had been noted earlier by Alberti, There are several possible ways Harvey could have come to
Canano, and Lusitanus) realize Fabricius was wrong about the function and purpose
3 There is a contrast between the venous system and the of the venous valves. One was through the influence of
arterial Salomon Alberti, as previously discussed. Another was the
4 This comparison points out the lack of physiological similarity of venous valves to the one-way heart valves. He
symmetry between veins and arteries. (In De motu cordis, knew this either from his own dissections, or from Alberti,
Harvey wrote: 'Let it be added that there are no valves Canano, and Lusitanus, or from Fabricius' pictures (see

Figure 4 Part of Table il from Hieronymus Fabricius' De venarum ostiolls. Figure ii improves on Alberti's picture (Figure 1 of this paper). It
shows the veins in an arm and two leg veins turned inside-out. The valves are cusps on the vein's wall. They are membranes which open
when blood flows against their edges, as depicted 495
 JOURNAL OF THE ROYAL SOCIETY OF MEDICINE Volume 88 September 1995

triple ostiola had been found in veins of oxen (similar to

those of the horse, as pictured in Figure 6). Yet, in his own
words, the triple cusps in the heart 'prevented reflux of
blood'. Obviously the door was open, so to speak, for
anyone to reason that the tricuspid venous valve could stop
blood flow in the ox's vein, just as the tricuspid cardiac valve
did in the heart. Even if the venous valves were not 100%
competent, their abundance effectively stopped reverse flow.
The above are reasons that could have led Harvey to
reject Galen's and Fabricius' explanations on the movement
of the blood. Any one of them, or a combination of them, or
all of them could have triggered Harvey's rejection.
..... ...............
Whatever the exact reasons, by 1617 Harvey had made
Figure 5 Two of the four figures from the only illustration in the conceptual leap. As the venous valves prevent the reflux
William Harvey's De motu cordis, 1628. With the text, they provide of the blood, he had to reject the old physiology. Yet,
a demonstration that the venous valves are one-way. Harvey's text Harvey had no immediate replacement physiology.
on this demonstration derives from Fabricius' De venarum ostiolis.
The muscular arm at the top is nearly a mirror image of that in Therefore, he was not ready to publicize his doubts.
Figure 4 of this paper Years later, when Boyle asked Harvey what first induced
him to think that the blood circulates, Harvey remembered
correctly. His answer in Disquisition, about the abundance
and orientation of the venous valves, expanded on one
statement from Prelectiones. Also in this statement is the link
between the structure of the venous and cardiac valves that
Boyle referred to in Considerations. Thus, both reports which
Boyle dictated about Harvey are supported by Harvey's early
lecture notes. This support reconciles Boyle's accounts of
Harvey's discovery: the keys to rejecting the current
paradigm were both the orientation of venous valves, their
abundance (which, in total, prevented reverse flow), and
*. - pjg. their similarity to the one-way heart valves (which did
sitI Pi. 2.

Figure 6 Fabricius had observed a tricuspid venous valve in an ox.

prevent reverse flow).
Pictured here are the multiple cusps in a horse's vein (from Figures Like Lusitanus, Canano, and Alberti, Harvey, at the time
1-8 Pettigrew, Trans R Soc Edin, 1863-1864, by kind permission of he wrote Prelectiones, understood the function of the valves,
the Royal Society of Edinburgh). (1)-(2) External jugular veins of but not their purpose. Using purpose in anatomy and
horse shown inside-out. Depicts valves, consisting of two (de),
three (abc), and four (fgh) cusps. (3) Section of external jugular physiology was part of Harvey's method and those before
vein of horse. Shows valve, consisting of two cusps (ab), with him. He followed 'the way of the anatomists'24 and would
dilations (g), corresponding to the sinuses of Valsalva in the not stop just knowing the function of the valves. He had to
arteries. (4) External jugular vein of horse opened. To show the
relation of the cusps (ab) above (re). (5) Portion of femoral vein
address their purpose, just as Fabricius had (incorrectly)
distended with plaster of Paris. Shows dilations (hg) in the course described the function and explained the purpose of these
of the vessel corresponding to the position of the valve. (6) Shows valves. Harvey's search for the purpose of the one-way
venous valve, consisting of two cusps (ab), in action. (7) The same, venous valves involved other 'investigations'.
not in action. (8) Venous valve from external jugular of horse,
consisting of three cusps

HARVEY'S HYPOTHESIS
Figures 2 and 3). Other reasons originate with Fabricius' De In Chapter 8 of De motu cordis, Harvey wrote how he
venarum ostiolis. First, was the anomalous orientation of the hypothesized that the blood circulates:
valves in the jugular veins, as Harvey noted in Chapter 13 of
De motu cordis. Secondly, Fabricius observed that ostiola In truth, when, from a variety of investigations through dissection of
could hold back venous blood and thought thick blood could the living in order to experiment and through the opening of arteries,
be held back for a long time. A third reason was from the symmetry and magnitude of the ventricles of the heart and
of the vessels entering and leaving (since Nature, who does nothing in
experimental. Fabricius pointed out that anyone pushing vain, would not have needlessly given these vessels such relatively
'the blood down through the veins would feel the resistance large size), from the skillful and careful craftsmanship of the valves
496 and power of the ostiola'. Finally, Fabricius reported that and fibres and the rest of the fabric of the heart, and from many other
 JOURNAL OF THE ROYAL SOCIETY OF MEDICINE Volume 88 September 1995

things, I had very often and seriously thought about, and had long When Harvey finally realized the implications of the
turned over in my mind, how great an amount there was, that is to blood's quantity, the conception of the blood's circulation
say how great the amount of transmitted blood would be [and] in how
I began
started to come into focus. Apparently, he then understood
short a time that transmission would be effected,
why the orientation of all the valves was opposed to the
. . .

privately to think that it might rather have a certain movement, as it

were, in a circle, . . .2 heart: the valves allowed circulation in one direction only.
The entire hypothesis finally became crystal clear to Harvey
While there is much in this passage, an important part is and he had his discovery. He now had to convince everyone
Harvey's list of a 'variety of investigations' which led to his else. Only during this later context of justification did
hypothesis. It includes: Harvey make his famous 'calculation' of the quantity of
blood, which was really more of a thought-experiment than
a calculation.
1 Vivisections and experiments
2 The symmetry and magnitude of the heart ventricles and CONCLUDING REMARKS
associated vessels
3 The skilful and careful craftsmanship of the heart valves Harvey sought to solve nature's mysteries. He dissected and
as well as other parts of the heart
vivisected. He studied the slowed-down heartbeats of dying
4 Many other things animals. He experimented with tourniquets both loosely and
5 The amount and transmission time of the blood tightly applied. He examined simple and compound hearts.
transmitted by the heart If Harvey had discovered the capillaries, then he would have
recognized an anomaly. However, this discovery was made
later, with an improved microscope. Instead, Harvey joined
Of interest in (2) listed above is the comparatively large his mass of accumulated data with the idea of purpose to
size of the heart vessels. The ventricles were discovered long reason that the orientation and abundance of venous valves
ago, and nothing appeared abnormal about them. and their similarity to the one-way cardiac valves demolished
Nevertheless, Harvey judged that they were too large for Fabricius' and Galen's physiology. Harvey had previously
their role in the current physiology. Since he thought that mentioned these keys to his discovery in Prelectiones (1616).
purposeful nature 'does nothing in vain', he had to find a After rejecting the old paradigm, Harvey sought the right
different explanation for this anomaly. explanation. He eventually hypothesized, perhaps as early as
For Harvey, it followed that the 'relatively large size' of 1617, that the blood circulates. Whatever the exact date, he
the ventricles and their conduits made it likely that blood certainly had written about his discovery by the middle of
was abundant in the body. However, it appears that this
1619.
abundance was a derivative factor from the prior Harvey made additions to his lecture notebook,
consideration of the conduits' large size. If the blood's Prelectiones, at various times up to at least 1626. One of
quantity initially led to the circulation hypothesis, then this these later additions is a short entry on the discovery of the
quantity would explain the vessels' large size. In which case, circulation. This addition is a main reason why many recent
Harvey would have said so-but he did not. Instead, he scholars think Harvey discovered the blood's circulation in a
wrote that:
later time frame. Yet Harvey did not record anything else
about the circulation or correct earlier erroneous statements
since Nature, who does nothing in vain, would have needlessly given based on the old physiology. His relative silence on an
these vessels such relatively large size.
important discovery especially perplexed the late Gweneth
Whitteridge. Concerning Prelectiones, she wondered:
The ventricles and their conduits have been designed large
and the question is for what purpose? If Harvey used this manuscript, as the additions to it imply, up to at
least 1626, we may well ask why it contains no references to the
Only now is Harvey ready to address the blood's crucial experiments by which the circulation was proved and why all
quantity. Quantity is important, but of the many factors statements which are inconsistent with the hypothesis of the
involved, the text provides no justification that it was prior. circulation are left uncorrected5.
Also, Harvey did not calculate amounts of blood. Rather, he
deduced from the comparatively large size of the vessels that Her line of questioning is based on her estimate that
an abundance of blood is involved. This abundance does Harvey's full discovery of the circulation was in 1625. This
become important and poses questions of its own. One is the article has shown that both the discovery of the circulation
purpose-based question, why is there so much blood passing and the composition of De motu cordis was earlier than
through the heart in so short a time? Thus it seems that the thought. That is why Harvey did not bother updating his
quantity of blood was one of the last factors Harvey Prelectiones. He was already writing De motu cordis and
considered. recording the crucial experiments and all else concerning the 497
 JOURNAL OF THE ROYAL SOCIETY OF -MEDICINE Volume 88 September 1995

discovery in it. Therefore, we should not be surprised that Basel: S Karger, 1976. Keynes G. The Life Of William Harvey. Oxford:
The Clarendon Press, 1966/1978
Harvey's lecture notebook contained only one additional
6 Harvey W. An Anatomical Disputation Concerning the Movement Of the
comment about his discovery. (He may have made this entry Heart and Blood In Living Creatures. Whitteridge G, transl. London:
prior to, and for, his January 1618 lecture.) Blackwell Scientific Publications, 1976
With his discovery of the blood's circulation, Harvey 7 Ferrario EV. William Harvey's debate with Caspar Hofmann on the
also became the discoverer of the real purpose of the venous circulation of the blood. J Hist Med 1960;15:7-21
valves. These valves now became important for 8 Bylebyl JJ. The growth of Harvey's De motu cordis. Bull Hist Med.
demonstrating that the blood circulated. Harvey only had 1973;47:427-70. Whitteridge G. Notes and comments. De motu cordis:
written in two stages? Bull Hist Med 1977;51:l130-9
to show that they were one-way. The experiments with the 9 Whitteridge G, transl. The Anatomical Lectures of William Harvey.
veins in a ligated arm (see Figure 5), were a convenient way Edinburgh: E & S Livingstone Ltd, 1964
to do this. Applying a tourniquet to the upper arm was the 10 Sanders JB, O'Malley CD. Andreas Vesalius Bruxellensis: the Bloodletting
first step in the common medical practice of bleeding a Letter of 1539. New York: Henry Schuman, 1947
patient. Anyone could then see the number of valves and, by 11 Adelmann HB. Introduction. The Embryological Treatises of Hieronymus
experiment, demonstrate that the function of the valves Fabricius of Aquapendente. New York: Cornell University Press, 1942
allows blood to flow only toward the heart. The next 12 Fabricius H. De venarum ostiolis (1603). Franklin KJ, transl. Springfield,
IL: CC Thomas, 1933
question is why?/for what purpose? The answer is that the
13 Pilkington R. Robert Boyle, Father of Chemistry. London: John Murray,
valves allowed one-way flow toward the heart because that 1959
was the direction the blood was moving in its circulation. 14 Lefanu WR. Notable Medical Books. Indianapolis: The Lilly Research
Thus, the orientation, one-way nature, and relatively large Laboratories, 1976
number of venous valves not only became important in 15 Hunter RA, Macalpine I. William Harvey and Robert Boyle. Notes Rec R
proving the hypothesis that the blood circulates, but also Soc Lond 1958;13:115-27
were keys to the initial discovery, as Harvey related to Boyle 16 Frank RG Jr. Harvey and the Oxford Physiologists. Los Angeles: The
University of California Press, 1980
years later.
17 Keynes G. A Bibliography Of the Writings of Dr William Harvey, 2nd edn.
Acknowledgments Part of the research for this paper was Cambridge: At the University Press, 1953
funded under a grant from The Pascal Centre, Redeemer 18 Boyle R. A Disquisition About the Final Causes Of Natural Things: Wherein it
is Inquired, Whether, And, (If At All) With What Cautions a Naturalist Should
College, Ancaster, Ontario. Opinions expressed in this Admit Them? London, 1688
article are not necessarily those of The Pascal Centre or 19 Whitteridge G. William Harvey and His Age [Book Review]. Isis
Redeemer College. I thank C Crouch and S McMullen for 1980;71 :503
their advice and comments. 20 French R. William Harvey's Natural Philosophy. New York, NY:
Cambridge University Press, 1994
21 Bylebyl J1. Boyle and Harvey on the valves in the veins. Bull Hist Med
REFERENCES 1982;56:351-67
1 Lord Cohen of Birkenhead. The germ of an idea, or what put Harvey on 22 Harvey W. Lectures On the Whole Of Anatomy. O'Malley CD, Poynter
the scent? J Hist Med Allied Sci 1957;12:102-5 FNL, Russell KF, transl. Berkeley: University of California Press, 1961
2 Banyon HP. The significance of the demonstration of the Harveian 23 Leibowitz JO. Early accounts of the valves in the veins. J Hist Med
circulation by experimental tests. Ann Sci 1938;3:435-56 1957;12:189-96. Franklin KJ. Valves in the veins: an historical survey.
3 Pagel W. William Harvey's Biological Ideas. New York: Hafner Publishing Proc R Soc Med 1927;21:1-33
Co., Inc., 1967. Bylebyl JJ, The medical side of Harvey's discovery: the 24 Wear A. William Harvey and the 'way of the anatomists'. Hist Sci
normal and the abnormal. In: Bylebyl Jj, ed. William Harvey and His Age. 1983;21:223-49
Baltimore: The Johns Hopkins University Press, 1979:28-102.
25 Bates DG. Harvey's account of his 'discovery'. Med Hist 1992;36:361-
4 Ghiselin M. William Harvey's methodology in De motu cordis from the 78
standpoint of comparative anatomy. Bull Hist Med 1966;40:314-27
5 Whitteridge G. William Harvey and the Circulation of the Blood. New York:
American Elsevier, Inc., 1971. Pagel W. New Light on William Harvey. (Accepted 27January 1995)

498